Professional Documents
Culture Documents
1999-2000
ISSN 1680-9270
September 2001, 2500 printed
Copyright lnternational Potato Center 2001
The lnternational Potato Center. 2001.
Scientist and Farmer Partners in Research for the 21 ST Century Program Report 1999-2000.
Lima, Peru. 480 p.
Contents
Program Overview
Wanda Collins, Deputy Director General for Research ................................................. 7
CIP's Research Project Portfolio ..................................................................................... 11
Building Partner Skills
P. Malagamba, E. Echeanda ....................................................................................... 16
Research on Potato
Plant Oefense Genes Associated with Quantitative Resistance to Potato Late Blight
P. Manosalva, S. Torres, F. Trognitz, R. Gysin, D. Nio-Liu, R. Simon,
M. Herrera, W. Perez, J. Landeo, B. Trognitz, M. Ghislain, and R. Nelson ................ 27
Characterization of Phytophthora lnfestans Colonizing Oifferent Solanaceous
Species in Peru, with lmplications on the Control of Potato Late Blight
G. Garry, G. Forbes, A. Salas, W. Prez, M. Santa Cruz, H.M. Pinedo,
E. Gonzles, M. Rivera, and R.J. Nelson ..................................................................... 39
Evaluation of Wild Potato Species for Resistance to Late Blight
W. Prez, A. Salas, R. Raymundi, Z. Huamn, R. Nelson and M. Bonierbale ........... 49
Quantifying Genetic Variance for Horizontal Resistance to Late Blight in
Potato Breeding Population B3C1
J.A. Landeo, M. Gastelo, G. Beltran, and L. Diaz ...................................................... 63
The Effect of Nitrogen Fertilization on Potato Late Blight in the Field
H.S. Jurez, J.R. Amaro, M.O. Rivera, A. Prraga, and R.J. Hijmans ......................... 69
Relationships of Fungicide Application to Late-Blight Oevelopment and
Potato Growth Parameters in the Tropical Highlands of Uganda and Kenya
O.M. Olanya, R. El-Bedewy, P.S. Ojiambo, P.T. Ewell, and J.J. Hakiza ..................... 77
Simulating Potato Late Blight in the Highland Tropics
G.A. Forbes, M.G. Chacn, M.V. Taipe, and R.J. Hijmans ......................................... 87
Oiscovery and Evaluation of a Valuable New Source of Resistance to
PLRV: Solanum tuberosum subsp. Andigena
E. Mihovilovich, L. Alarcn, A.L. Prez, and M. Bonierbale ...................................... 95
Assessment of Latent lnfection Frequency in Progeny Tubers of Advanced
Potato Clones Resistant to Bacterial Wilt: A New Selection Criterion
S. Priou, C. Salas, F. de Mendiburu, P. Aley, and L. Gutarra ..................................... 105
Evaluation of Bt-cry1/a1 (cryV) Transgenic Potatoes on Two Species of
Potato Tuber Moth, Phthorimaea opercu/ella and Symmetrischema
tangolias (Lepidoptera: Gelechiidae) in Peru
A. Lagnaoui, V. Caedo, and D.S. Douches ............................................................. 117
Assessment of the lnactivation Time of Phthorimaea opercu/ella Granulovirus
(PoGV) at Oifferent lntensities of Natural lrradiation
M. Sporleder, O. Zegarra, J. Kroschel, J. Huber, and A. Lagnaoui ............................ 123
lntegrated Control of Potato Bacteria! Wilt in Kabale Oistrict, Southwestern
Uganda
B. Lemaga .................................................................................................................. 1 29
Optimization of Sample Size for the Oetection of Latent lnfection by Ralstonia
solanacearum in Potato Seed Tubers in the Highlands of Peru and Bolivia
S. Priou, R. Torres, A. Villar, J. Martinez, O. Barrea, L. Gutarra, and
F. de Mendiburu ......................................................................................................... 143
CIP Program Report 1999-2000
Research on Sweetpotato
Transgene Expression of Rice Cysteine Proteinase lnhibitors for the Development
of Resistance against Sweetpotato Feathery Mottle Virus
G. Cipriani, S. Fuentes, V. Bello, L.F. Salazar, M. Ghislain, and O.P. Zhang ........... 267
AFLP Markers
A. Kriegner, J.C. Cervantes, K. Burg, R.O. Mwanga, and O.P. Zhang ....................... 303
Program Overview
Scientist and Farmer. Partners in Research far the 2 7st
Century. The title of the 1999-2000 Program Report
exemplifies not only the scope of CIP's research, butthe
philosophy behind it. As a Future Harvest Center (see
note, page 9), CIP's overarching goals are to reduce
poverty, ensure food security, and protect natural resources. Reaching those objectives depends on
developing and testing new technologies in the lab and
in the field. However, it is only after successful application and adoption in the fields of resource-poor farmers
around the world that the benefits of those technologies
can be seen and felt. Scientist and farmer partnerships
are helping us to accelerate the adoption and application of new technologies.
We learn then we
share Modern global
Broad-based Research
During the two years of research included in this Program Report, CIP carried out its program through 17
research projects addressing the most pressing needs of
developing country farmers (see page 11 for project
overviews). With a mandate encompassing potatoes,
sweetpotatoes, Andean root and tuber crops, and natural resource management, CIP has much to report.
In the fol lowing pages, there are research reports ranging
from the application of highly complex molecular
technologies to identify plant defense genes for late
blight resistance, to the development and evaluation of
information systems
(GIS) use sophisticated techniques that
link satellite information and computer
models with many
research disciplines.
A series of research
projects reported here
show the practica!
side of this type of
work. Qui roz et al.
(page 3 61 ) report on
how GIS can be used
to define agricultura!
zones in the
Cajamarca, Peru,
watershed. Baigorria
et al. (page 371) write
about weather in that
mountain environment and Bussink and
Hijmans take a look at
land use change there
from 1975-96 (page
421 ). Finally, Posner
et al. (page 413)
present the practica!
application of this
combined knowledge
when they report on
CONDESAN-in itiated
soil conservation
planning in two
communities in the
same watershed.
Researchers, NGO
workers, and commun ity leaders worked
together to consider
land use options.
Program OVerview
Wanda Collins
Deputy Director General for Research
Teaching farmers
from ali walks of life
"The project [FFS in
Nepal] demonstrated
that one does not
have to be amale,
1iterate farmer from
the dominant ethnic
group to participate
meaningfully in
agricu ltu ral research
and extension. Of
those who successfu 1ly completed the
FFS, one out of four
was i 11 iterate, one out
of three was female. In
addition, 7 of the 9
FFSs had participants
coming from multiple
ethnic groups.
Average knowledge
gain on patato seed
and disease management was 84%,"
report Hidalgo,
Campilan, and Lama.
(See page 239).
Future Harvest builds awareness and support for food and environmental research for a world with
less poverty, a healthier human family, well-nourished children, anda better environment. Future
Harvest supports research, promotes partnerships, and sponsors projects that bring the results of
research to rural communities, farmers and families in Africa, Latin America, and Asia. lt receives its
principal funding from a group of governments, private foundations, and the CGIAR.
CIP's Directors
Director General, Hubert Zandstra
Deputy Director General for Research, Wanda Collins
Deputy Director General for Finance/Administration, Hector Hugo Li Pun
Director for lnternational Cooperation, Roger Cortbaoui
15
16
17
2
3
4
5
6
7
8
9
10
11
12
13
14
genetic resources
1O
Leader
R. Nelson
S. Priou
L. Salazar
A. Lagnaoui
M. Bonierbale
E. Chujoy
T. Walker
L. Salazar
E. van de Fliert
O.P. Zhang
O.P. Zhang
G. Prain
T. Walker
R. Quiroz
W. Roca
M. Hermann
M. Holle
Project 2
lntegrated control of bacteria! wilt
(S. Priou)
Bacteria! wilt (BW), caused by Ralstonia
solanacearum, is the second main potato
disease in the world. The key to overcoming the constraint is to control the spread
of the disease by using only healthy
planting material. CIP's research therefore
concentrates on developing tools to detect
the presence of bacteria in soil and tubers.
This knowledge is used to improve crop
management and seed systems and to
identify available resistant/tolerant
material. The project also aims at designing, validating, and promoting integrated
strategies for managing bacteria! wilt in
different production systems and complet-
Project 3
Control of potato viruses
(L. Salazar)
Virus diseases cause serious losses in
potato and also disrupt global efforts to
improve potato, because national and
inter-national regulations control the
movement of virus-infected seed and
genetic resources. Biotechnology tools are
used to identify resistance genes in related
plant species and to clone and transfer
them to potato; these tools are combined
with traditional methods of breeding to
develop adapted cultivars resistant to a
range of viruses. ldentifying and characterizing the most important viruses and
virus-like agents that affect potato are
essential steps toward developing sensitive, low-cost methods for large-scale
detection. CIP researchers also study
epidemiological factors that affect virus
spread, with particular attention to interaction between viruses and other pathogens
that may affect plant resistance response,
and train national scientists in virus
identification, detection, and control
techniques. Research and training activities focus on the most important potato
viruses (PLRV and PVY) and on the potato
spindle tuber viroid (PSTVd). Particular
attention is given to practica! utilization/
adoption of virus-resistant materials
already produced at CIP (including
genotypes already carrying combined
resistance to more than one virus, preferably in a multiplex condition).
11
Project 4
Project 6
(A. Lagnaoui)
Key pests of potato, globally or regionally,
are three species of potato tuber moth (a
threat that is rapidly becoming more
serious), severa! species of Andean potato
weevil, the leaf-miner fly, whiteflies and
severa! flea beetles. Nematodes that
reduce potato yields and favor the development of bacteria! wilt pathogens are
potato cyst, rosary, and root-knot nematodes. This project seeks to develop
locally-adapted integrated pest management programs for these pests,
emphasizing sustainable, ecologicallybased, and economically-sound practices
that will lead to reduced use of chemical
pesticides and increased benefits for
farmers. Components include biological,
cultural, and resistance aspects of control.
E. Chujoy)
True potato seed (TPS) enables a crop to be
grown in areas where traditional production systems fail, for example, where seed
tubers are scarce or not available. By
facilitating the transfer of TPS technology
in such areas of the tropics and subtropics,
this project aims to expand patato cultivation and increase its efficiency (reduce
production costs, increase yields). CIP
concentrates on improving parents for
hybrid TPS production and developing
needed specific traits such as late-blight
resistance, earliness, and seed set. CIP's
work is backstopped by local organizations (private sector, NGOs, NARS) in
efforts to commercial ize TPS systems and
thus underpin developing small industries.
Project 5
Project 7
/ M. Bonierbale)
In many countries, the lack of efficient
formal and informal seed potato systems
has limited the diffusion of new and
improved varieties because only limited
amounts of healthy clona! planting
material are available. Varietal introduction and diffusion is dependent on the
informal system, but it must be linked with
the formal system and it must emphasize
high quality planting material. This project
provides research and technical assistance
to selected formal and informal seed
systems in various countries to help them
improve their efficiency and effectiveness.
This is accomplished through farmer
training and establishment of pilot seed
systems. The project also explores innovations in linkages between formal and
informal seed systems, aiming to speed up
varietal introduction and diffusion.
12
Project 8
Control of sweetpotato viruses
(L. Salazar)
Project 9
lntegrated management of sweetpotato
pests (E. van de Fliert)
The aim of this project is to develop IPM
(integrated pest management) systems for
sweetpotato. These systems need to be
compatible with farmers' crop management practices, as well as with prevailing
ecological and socioeconomic conditions,
to ensure effective and sustainable solutions. Therefore, participatory approaches
are applied to prioritize research needs,
develop adapted pest and crop management components, and design learning
strategies which enhance farmers' ecological knowledge and decision-making and
Project 1 O
Postharvest utilization of sweetpotato
(G. Scott (until August 2000)/D.P. Zhang)
This project studies technologies to
improve the livelihoods of rural poor
through diversification and expansion of
sweetpotato use. The main beneficiaries
are women and children and small households. Nutrition and income are improved
and poverty is reduced. Project goals
include facilitating the development of
small enterprises based on added-value
from primary processing (e.g., starch and
flour), and the more efficient use of
sweetpotato roots, vines, and by-products
as animal feed. In Africa, the goal is to
enhance food security by taking advantage of sweetpotato's nutritional qualities.
CIP researchers evaluate opportunities and
undertake collaborative research on
markets, raw material quality, process
development, product qual ity, and the
social acceptability of innovation in pilot
enterprises. They tap such resources as
NARS, NGOs and users in target countries,
along with global centers of research
excellence in disciplines not available inhouse, such as food science/technology
and animal sciences.
Project 11
Breeding sweetpotato for high-dry-matter
yield and adaptation (D.P. Zhang)
This project aims to improve sweetpotato
production and use through the development and adoption of high-dry-matter/
high-starch varieties with adaptabil ity to
low-input, subsistence farming systems.
The diverse sweetpotato germplasm at CIP
13
Project 12
Strategic lnitiative on Urban and Periurban Agriculture (G. Prain)
The Strategic lnitiative on Urban and Periurban Agriculture (SIUPA) was launched
by the CGIAR in late 1999 in response to
growing urbanization and increased
dependence of city dwellers on farming:
CIP is the convening center for the initiative. SIUPA's goals are to contribute to
increased food security, improved nutritional status, and higher incomes of urban
and peri-urban farmers; reduce the negative environmental impact of urban and
peri-urban agriculture and enhance its
positive ecological potential; and establish
the perception of urban and peri-urban
agriculture as a positive, productive, and
essential component of sustainable cities.
In collaboration with the many national
and international efforts that have started
in recent years to address the issue of
urban and peri-urban agriculture, SIUPA is
establishing, in regional sites, a set of
research activities collectively known as
Urban Harvest. Several international
agricultura! research centers are already
working in such areas as technology and
policy aspects of urban agriculture and
14
Project 13
Sustainability of rice-based cropping
systems featuring potato as a cash crop
(T. Walker)
In response to increasing land scarcity in
subtropical South and Southeast Asia,
potatoes with a high production potential
per unit time are increasingly being
planted in intensive sequential cropping
systems, such as rice-patato-rice, and in
more intensive intercrops, such as ricepotato/maize. Typically, rice is planted at
the onset of the rainy season, irrigated
patato is sown in the cooler dry season,
and irrigated rice or another crop is
cultivated in the hot summer. To realize
the potential of the patato crop, factors
that threaten the sustainability of these
input-responsive cropping systems must be
identified and addressed. This project
diagnoses constraints to increasing and
maintaining productivity in selected
potato and rice-based cropping systems;
and generates crop and natural resource
management information on how to
alleviate the most important of those
constrai nts.
Project 14
Sustainable land use in the Andes
(R. Quiroz)
The Andes comprises a series of unique
habitats rich in natural resources. The
inhabitants of this region confront massive
poverty, increasing population growth, and
rapid degradation of the natural resource
base. They face the difficult challenge of
trying to increase agricultura! productivity
while simultaneously decresing stress on
Project 17
Conservation and characterization of
Andean root and tuber crops (M. Halle)
Project 15
Conservation and characterization of
potato genetic resources
(Z. Huamn (until June 2000)/W. Roca
CIP holds the most comprehensive collection of germplasm of wild and cultivated
potatoes in the world. Key objectives of
this project include safe, long-term
conservation and characterization of the
germplasm, and developing a database
containing all information on the collection and making this information available
to interested parties. Research attempts to
in crease uti 1ization of the pota to genetic
diversity by identifying key desirable traits
and distributing healthy seed stocks and
clonal materials throughout the world for
use in patato improvement programs. The
project provides key input into CIP's own
breeding efforts.
Project 16
Conservation and characterization of
sweetpotato genetic resources
(M. Hermann)
Assisting national programs in rationalizing strategies for both ex situ and in situ
conservation of Andean root and tuber
crops (ARTC) involves the study and
preservation of biodiversity, with emphasis
on four priority genera - Oxalis (ocat
Ullucus (ulluco), Canna (achira), and
Arracacia (arracacha), including wild
species-and on the material of Mirabilis
expansa (mauka), Pachyrhizus ahipa
(ahipa), Smallanthus sonchifolius (yacon),
Tropaeolum tuberosum (mashua), and
Lepidium spp (maca). The project is
systematically assessing the potential of
ARTC to promote wider use in the subtropical and tropical highlands, within and
outside the Andean region, through the
study of current marketing and consumption patterns. lt aims to identify latent
demands these crops may satisfy in the
future, and to produce healthy planting
materials for farmers. This project is
perhaps the only one in the world with a
majar effort towards developing virus
identification and eradication procedures
for these important, underutilized crops.
15
Training at Headquarters
Individual
A large part of individualized training took
place on-site in CIP's labs and greenhouses
16
Group training
Group courses cover all CIP's research
areas and concentrate on transferring
methodologies (Table 2, Figure 3), whereas
workshops focus on planning, information
exchange, and discussions on the progress
of ongoing collaborative activities. A
popular group training topic in 1999-2000
was biodiversity in relation to Andean root
and tuber crops. Several workshops were
held on integrated management of the
most destructive pests in potato and
sweetpotato. Others were held to facilitate
progress for activities supported by
FONTAGRO, (funded by IDB), IFAD,
DFID, UNEP, IDRC, SDC, and other
organizations. A number of courses and
workshops were organized in collaboration
with our regional headquarters' staffs and
held away from CIP headquarters.
As our partners improve their skills they
become future col laborators and contact
points in their home country, thereby
D
17%
39%
12%
17%
20%
17%
D
D
17%
11 %
17
of Agricultura! Sciences
Canary lslands,
Spain
Table 1. (continued)
Molecular techniques applied to patato Peru (2)
characterization
Molecular techniques applied to patato Spain
genetic diversity
Molecular techniques applied to
Peru (3)
germplasm characterization
19
11
15
19
7
Workshop/Ecuador
20
Meeting/Peru
Conference/Ecuador
20
163
CourseNietnam
Course/Kenya
Workshop/I ndia
Workshop/China
Workshop/China
Workshop/Peru
Course/Peru
Course/Cuba
Course/Peru
Workshop/Andean
countries
Course/Egypt
Course/Kenya
12
10
33
20
48
180
6
11
27
15
Bangladesh, India
China
Peru
Argentina, Bolivia, Ecuador, Mexico,
Peru
Bolivia, Colombia, Ecuador, Peru,
Venezuela
Peru
Argentina, Austria, Bangladesh,
Belarus, Bolivia, Brasil, Cameroon,
China, Colombia, Cuba, Costa Rica,
Czech Republic, Denmark, Ecuador,
England, France, Germany,
Honduras, India, Israel, Kenya,
Korea, Mauritius, Mexico,
Netherlands, Nepal, Norway,
Panama, Peru, Polony, Spain,
Sweden, Switzerland, Uganda,
Uruguay, USA, Venezuela
Vietnam
Kenya
India
China, Philippines, Peru
China
Peru
Bolivia
Cuba
Peru
Andean countries
20
6
Egypt
Burundi, Ethiopia, Cameroon,
Kenya, Uganda
Course/Kenya
Workshop/Bangladesh
9
15
Workshop/I ndonesia
23
20
Co-sponsors1
CIP/Association far Andean
Technical-Cultural Promotion,
(Peru)/lnter-American Foundation
(USA)
Germany-1 mprov. ResisVUPWARD
lnternational Fund far Agricultura!
Development
FORTIPAPA, INIAP
Global lnitiative on Late Blight
Global lnitiative on Late Blight
UPWARD
Programme Rgional de
l'Amlioration de la Culture de la
Pomme de Terre et de la Patate
Douce en Afrique Central et de l'Est
(CIP network)
PRAPACE
lnternational Fund far Agricultura!
Development
Users' Perspective with Agricultura!
Research and Development (CIP
network)/Swiss Development
Cooperation
Table 2. (continued)
Workshop/China
WorkshopNietnam
47
20
China
Indonesia, Vietnam
Course/Ecuador
Course/Ecuador
Course/Peru
Cou rse/Swaziland
22
39
4
22
35
WorkshopNietnam
Course/Bangladesh
Workshop/Nepal
Workshop/Nepal
Workshop/Peru
85
Workshop/Peru
34
Course/Peru
21
11
16
12
Workshop/Kenya
15
Workshop/lndia
11
WorkshopNietnam
15
Postharvest quality, nutrition,
Workshop/Peru
18
Course/Kenya
11
Workshop/Kenya
31
Workshop/Cuba
16
Workshop/Peru
31
Workshop/Malawi
27
21
Table 2. (continued)
Activity/venue
Part. (#)
Workshop/Malawi
15
Workshop/Kenya
22
Co-sponsors1
42
Workshop/Peru
40
Workshop/Peru
Workshop/Colombia
8
18
WorkshopNietnam
30
Other
Workshop/Peru
32
Meeting/Ch in a
Workshop/Peru
Course/Brazil
50
12
36
Meeting/Peru
23
Meeting/Uganda
40
Meeting/Malawi
19
Course/Peru
31
Course/United
Kingdom
37
Course/Peru
25
22
Biodiversity Project
Chinese Academy of Agric. Sci.
Table 2. (continued)
Workshop/Brazil
Workshop/Ethiopia
Workshop/Ethiopia
Total
44
15
5
Brazil, Ecuador
Ethiopia
Burundi, Ethiopia, Kenya, Uganda
FAPESP/EMBRAPNCENA
lnternational Fund far Agricultura!
Development
1DRC/PRAPACE
2140
1 Where
23
270
1371
175
383
2599
Bolivia
China, Uganda,
Ethiopia,
Bangladesh
Ethiopia
Peru
LB = late blight; BW = bacteria! wilt; PTM = patato tuber moth; APW = Andean patato weevil.PROINPA = Fundacin
para la Promocin e Investigacin de Productos Andinos, Bolivia; ASAR - Asociacin de Servicios Artesanales y Rurales
de Bolivia.
24
Research on Potato
Scientist and Farrner
Partners in Research for the 21 st Century
27
28
Research on Potato
DNA markers
The following ONA clones were used for
analysis of restriction fragment length
polymorphism (RFLP): Pto, Pti-4, Pti-5, and
Pti-6 (provided by M. O' Ascenzo and
G.B. Martin), glucanase (provided by
E. Kombrink), osmotin (provided by
T. Chen), PAL and 4CL (provided by
l.E. Somssisch), ch aleone isomerase (CH 1)
from maize (provided by E. Grotewold),
phosphol ipase O probe (provided by
O. McGee and J.E. Leach), SGT (provided
by W. Belknap), and 21 anonymous
probes previously associated with QTLs
for late-blight resistance (provided by
C. Gebhardt).
For PCR-based assays, gene sequences and
sequences of expressed s,equence tags
(EST) were obtained from the Mendel
database and the database of the National
Center for Biotechnology lnformation
(NCBI). Corresponding primers were
designed using the program Primer 0.5
(unpublished software by Oaly, Lincoln,
and Lander; available from the Whitehead
lnstitute, MIT Center for Genome Research) and synthesized by GENSET (La
Jol la, CA, USA).
Data analysis
lndependence of marker distribution
(presence or absence) and phenotypic
groups (resistant and susceptible) was
tested by the X2-test using a 2x2 contingency with one degree of freedom. To
reduce the experiment-wise error, we used
a maximum P value of 0.01 to declare a
significant association. MAPMAKER/EXP
v3.0b software was used to locate the
candidate genes on the PO map (Lander
and Botstein, 1989).
<o.os.
Results
In this study, trait-marker associations were
sougrt for quantitative resistance to late
blight using genes involved in plant
defense. A total of 335 segregating loci
were assayed in two populations of patato,
segregating for quantitative resistance to
late blight. The loci tested in this study
represented 39 genes, gene elements, or
gene families, many of which are known
or suspected to play a role in plant defense. Loci were assayed using four
methods: ONA hybridization (RFLP),
simple PCR, CAPS, and LM-PCR (Hornstra
and Yang, 1993). The results were compared to existing datasets obtained using
anonymous markers.
29
w
e
:o
:::i
"'O
o~
Table 1. Loci associated with quantitative resistance to late blight in a diploid mapping population and in a tetraploid breeding population at P < 0.01.
x2
Target gene ormarker
Marker type Locus defined by (ID/enzyme/ Band origin
Number of observad (tested)
(parent)
/band) molecular weight (kb)
r
s
Resistance genes (pathogen recognition)
p
3 (16)
Conserved domain
PCR
12 (17)
AS 1-S2/0.23
6.96
Signal transduction and gene regulation
Pti-5
o
RFLP
3 (23)
Pti5/EcoRl/3.8
14 (23)
9.33
LM-PCR
o
WRKY
WRKY1-r/BamHl/0.33, 0.51
25 (34)
4 (33)
23.28
LM-PCR
o
WRKY2-r/BamHl/0.49
10 (34)
24 (33)
10.89
LM-PCR
o
WRKY2-r/BamHl/0.9
25 (34)
20.78
5 (33)
LM-PCR
o
18.62
WRKY4-f/K~nl/0.9
5 {34}
23 {33}
Pathogenesis-related genes
1,3-B-glucanase (GLU)
CAPS
387170.9
12 (26)
26 (29)
GLU/Dral/0.12
10.19
LM-PCR
o
Osmotin (PR-5)
Osmotin/EcoRl/0.29
26 (30)
5 (26)
22.97
o
RFLP
Osmotin/EcoRV/13, 7; Xbal/12
24 (33)
10 (33)
10.25
o
CAPS
Osmotin/Rsal/4
9 (31)
26 (34)
12.83
STH2-21 (PR-10)
CAPS
o
STH/Dral/0.65,Taq/0.45
5 (33)
16 (31)
8.06
o
CAPS
STH/Dral/0.3, 0.26, 0.6
5 (33)
14 (30)
5.99
CAPS
STH/Taq 1/0 .850
387170.9
16 (23)
6 (22)
6.44
CAPS
386209.10
STH/Taq 1/0 .259
21 (30)
3 (15)
8.14
CAPS
386209.10
18.02
STH/Dral/0.17
25 {33}
2 {19}
Phenylpropanoid pathway (phytoalexin synthesis and lignin synthesis)
p
Phenylalanine ammoniumlyase (PAL) RFLP
12 (34)
24 (33)
PAL/EcoRl/1.9
7.99
o
RFLP
10 (34)
23 (33)
9.32
PAL/EcoRl/2
o
RFLP
25 (34)
3 (31)
24.42
PAL/EcoRl/2.8
p
31 (34)
1o(33)
23.63
RFLP
PAL/EcoRl/3
o
RFLP
PAL/Hindlll/2.8, 2.7
9 (34)
22 (27)
16.09
p
15 (28)
4 (26)
7.027
RFLP
PAL/Hindlll/5
o
8 (28)
21 (26)
12.74
RFLP
PAL/Hindlll/5.5
o
26 (34)
16.24
RFLP
8 (33)
PAL/Ndel/3.6
o
9 (34)
27 (33)
RFLP
PAL/Ndel/4,4.4
18.47
p
RFLP
22 (34)
6 (33)
13.04
PAL/Ndel/5.9
o
RFLP
26 (34)
PAL/Ndel/6
1 (33)
34.55
o
RFLP
PAL/Ndel/6.8, 6.9
26 (34)
5 (33)
22.92
LM-PCR
o
PAL2/BamHl/0.075
9 (34)
26 (32)
17.72
P (x2 test)
0.008
0.002
<0.00001
0.00096
<0.00001
0.00002
0.001
<0.00001
0.001
0.0003
0.0045
0.014
0.011
0.004
<0.00001
>
0.004
0.002
<0.00001
<0.00001
<0.0001
0.008
0.0003
<0.0001
<0.00001
0.0003
<0.00001
<0.00001
<0.0001
Lipid metabolism
Lipoxygenase
Other
Metallo-thionein (Meth IA)
NADH-Deh~drogenase
LM-PCR
LM-PCR
LM-PCR
RFLP
RFLP
RFLP
LM-PCR
LM-PCR
PAL2/BamHl/0.41 O
PAL6/BamHl/O .420
PAL6a/BamHl/0.5
4CL/EcoRl/9
4CL/Hindlll/8
4CL/Ndel/4.5
CHl/BamHl/0.320, 0.315
CHl/BamHl/0.53
CAPS
D
D
D
D
D
D
D
D
27 (34)
23 (30)
9 (30)
24 (33)
25 (34)
9 (34)
27 (34)
8 (34)
6 (32)
4 (27)
22 (27)
11 (32)
12 (33)
22 (33)
5 (32)
24 (32)
21.89
19.39
13.18
8.13
7.9
9.33
24.36
15.48
<0.00001
<0.00001
0.0003
0.004
0.005
0.002
<0.00001
<0.0001
LOX/Rsal/0.28
386209.10
23 (28)
4 (20}
15.86
<0.0001
PCR
PCR
Meth IA
NADH dhase
D
p
4 (22)
17 {22)
16 (23)
5 {23)
10.03
11.74
0.002
0.0006
RFLP
RFLP
RFLP
RFLP
Cl21/EcoRV/6.5
CP116/Dral/3
GP186/EcoRl/8
GP313/Xbal/11
D
D
D
D
18 (30)
19 (25)
3 (25)
9 (29)
5 (31)
1 (23)
11 (21)
20 (29)
10.69
22.44
6.98
6.89
0.001
<0.00001
0.0082
0.0086
Anonymous markers
C')
=ti
""tl
ce
;
3
~
o
~
<O
<O
<O
1
N>
o
o
o
PAL
PO
p = 0.006
PO
WRKY
PO
P=0.0004
Figure 1: Ligation-mediated PCR of PAL and WRKY genes, for groups of progeny of the PD population with
contrasting phenotypes for late blight resistance under field conditions. "R" and "S" refer to the resistant and
susceptible groups respectively; "P" and "D" refer to the Solanum phureja and S. tuberosum dihaploid parents
respectively. The significance of linkage disequilibrium of the band is indicated by the probability value (P) of the
x2 test.
SSR markers were associated with resistance at a very low rate (0-5%).
Defense-related genes were associated
with resistance at higher frequencies
(5-46% for different groups of genes).
Map locations were estimated for many of
the resistance-associated markers identi-
32
Research on Potato
Table 2. Summary of results obtained far the candidate gene analysis of the PO and 393228 populations.
Loci associated with resistance by the chi-sguare test at P < 0.01 are considered as "hits".
Group
Assays
Segregating loci ResistanceHits per
Hits per
Gene family
(no.)
(no.}
assay (%)
associated loci
segregating
("hits") (no.}
locus (%}
Far candidate gene study
Overall
139
335
48
35
14
PO population (2x)
104
289
34
12
33
393228 (4x}
35
46
5
14
11
For candidate genes, PO population
R-genes
13
76
8
Signal transduction
5
44
21
23
11
and gene regulation
WRKY
11
4
133
3
36
Phenylpropanoid
21
12
175
63
33
pathway
PAL (RFLP)
26
3
12
400
46
PAL (RFLP +
38
16
42
6
267
LM-PCR)
PR-genes
38
5
22
13
16
o
Lipid metabolism
13
o
o
12
55
2
4
Other genes (PCR}
22
9
Candidate genes, 393228 population
PR-genes
11
22
4
36
18
1
Lipid metabolism
6
7
17
14
17
o
o
o
18
Other (PCR}
By method, candidate gene study (both populations}
106
3
10
3
PCR (direct)
30
8
62
15
13
CAPS
53
17
RFLP
99
50
17
34
68
11
50
16
LM-PCR
22
Data obtained for QTL mapping study, PO population
456
42
AFLP
43
98
9
7
216
16
14
RAPO
111
4
11
5
RFLP-genomic
78
38
2
13
6
32
SSR-genomic
16
o
o
o
RFLP-genes
28
14
104
5
10
5
SSR-genes
52
and most consistent QTL detected for the
PD population (Ghislain et al., 2001 ).
Using DNA hybridization and CAPS, no
osmotin bands were found for the QTL on
chromosome XII, but an osmotin-like band
associated with resistance was detected
and located to a peak of chromosome XII
using LM-PCR. WRKY and osmotin were
co-localized with QTL on chromosome
VIII. Sorne QTLs identified by Ghislain et
al. (2001) were not associated with any of
the markers identified in this study.
33
1 Quantitative trait loci for late blight resistance defined in the PO mapping population (Ghislain et al., 2001 ). Codes
(Q-x-y) indicate the following: Q refers to QTL; x refers to the parent from which the OTL was derived (P for Solanum
phureja; O for S. tuberosum dihaploid); y refers to the chromosome on which the QTL was located.
2 Only markers showing statistically significant association with resistance were considered.
using CAPS (n
PCR (n = 1).
Discussion
34
Research on Potato
= 1),
RFLP (n
= 1),
and LM-
Acknowledgments
This research was supported in part by the
government of Germany (BMZ Project No.
96.7860.8-001 .00). We are grateful to the
individuals and institutions that provided
ONA clones. We thank Ornar Prado for
technical assistance in the laboratory. We
35
References
Botstein, D., R.L. White, M. Skolnich, and
R.W. Davis. 1980. Construction of a
genetic linkage map of man using
restriction fragment length
polymorphisms. American Journal of
Human Genetics 32:314-331.
Constabel, C.P. and N. Brisson. 1995.
Stigma- and vascular-specific expression
of the PR-1 Oa gene of potato: A novel
pattern of expression of a pathogenesisrelated gene. Molecular Plant-Microbe
lnteractions 8(1):104-113.
Dellagi, A., J. Helibronn, A.O. Avrova,
M. Montesano, E.T. Palva, H.E. Stewart,
l.K. Toth, O.E. Coke, G.D. Lyon, and
P.R. Birch. 2000. A potato gene
encoding a WRKY-like transcription
factor is induced in interactions with
Erwinia carotovora subsp. atroseptica
and Phytophthora infestans and is
correlated with class 1 endochitinase
expression. Molecular Plant Microbe
lnteractions 13:1092-1101.
Eulgem, T., P.J. Rushton, S. Robatzek,
and 1.E. Somssich. 2000. The WRKY
superfamily of plant transcription
factors. Trends in Plant Science
5:199-206.
Ewing, E.E., l. Simko, C.D. Smart,
M.W. Bonierbale, E.S.G. Mizubuti,
G.D. May, and W.E. Fry. 2000. Genetic
mapping from field tests of quantitative
and qualitative resistance to
Phytophthora infestans in a population
derived from Solanum tuberosum and
Solanum berthaultii. Molecular Breeding
6:25-36.
.
Faris, J.D., W.L. Li, D.J. Liu, P.D. Chen,
and B.S. Gill. 1999. Candidate gene
analysis of quantitative disease
resistance in wheat. Theoretical Applied
Genetics 98:219-225.
Fritzemeier, K., C. Cretin, E. Kombrink,
F. Rohwer, J. Taylor, D. Scheel, and
36
Research on Potato
5:399-415.
Porebski, S., L.G. Bailey, and B.R. Baum.
1997. Modification of a CTAB DNA
extraction protocol for plants containing
high polysaccharide and polyphenol
2:225-238.
Shen, K.A., B.C. Meyers, M.N. lslamFaridi, D.B. Chin, D.M. Stelly, and
R.W. Michelmore. 1998. Resistance
gene candidates identified by PCR with
degenerate oligonucliotide primer map
to clusters of resistance genes in
lettuce. Molecular Plant-Microbe
lnteractions 11 :815-823.
Trognitz, B. R., M. Orrillo, L. Portal,
C. Romn, P. Ramn, S. Perez, and
G. Chacn. 2001. Evaluation and
analysis of reduction of late blight
disease in a diploid potato progeny.
Plant Pathology 50(3):281-291.
37
To test the hypothesis that wild and cultivated Solanum spp. are attacked by
the same populations of Phytophthora infestans (Mont.) de Bary, we characterized isolates attacking wild relatives of potato (Solanum tuberosum L.)
using molecular markers and by an aggressiveness parameter: diameter of
lesion in a detached-leaf assay. lsolates (n = 287) were sampled from the
northern and central highlands of Peru, from different cultivated potatoes
(both bred varieties and native potatoes), from different species of wild
potatoes of Petota section, from S. caripense (non-tuber bearing Solanum of
Basarthrum section), and from wild tomatoes. All the isolates analyzed belonged to four lineages, EC-1, PE-3, PE-7, or US-1, which previously had been
described in Peru. The same pathogen genotypes can attack wild (Petota) and
cultivated potatoes. Our results also showed sorne evidence of host differentiation. The genotype PE-7, mainly obtained from wild tomatoes, hada novel
peptidase genotype. A significant interaction (isolate origin) x (inoculated
host) for lesion diameter was obtained between isolates from S. caripense and
bred potatoes of the EC-1 or US-1 lineages, but not for other host-isolate
combinations.
Late blight (LB), caused by the oomycete
pathogen P. infestans is one of the most
devastating diseases of cultivated potatoes
worldwide. P. infestans can cause symptoms on leaves, stems, and tubers. The
disease is responsible for important
economic losses and high levels of
fungicide use. Because qualitative resistance conditioned by major genes has
proven to be non-durable, potato improvement efforts in both developing and
industrialized countries have focused on
improving the levels of quantitative
1
39
40
Research on Patato
Table 1. lsolates obtained from different wild Solanaceae and cultivated potatoes between 1997
and 2000, Peru.
Year collected
lsolates per department
Host
Ancash Cajamarca Huancavelica Junn Libertad Lima Piura
Cultivated (bred) potatoes
(113 isolates)
2000
TxA cv Amarilis-INIA
4
5
10
1999-2000
6
TxA cv Canchn
5
4
8
8
TxA cv Mix
2000
5
TxA cv Molinera
2000
2
1999-2000
5
15
TxA cv Perricholi
6
1999-2000
10
4
TxA cv Yungay
9
6
Cultivated (native) potatoes
(42 isolates)
4
4
S. tuberosum ssp. andigena
19991-2000
10
S. goniocalyx
1999-2000
5
7
1
1999-2000
1
S. choucha cv. Huayru
2000
1
6
S. phureja
2000
2
S. stenotonum
Wild potatoes (70 isolates)
1998
S. acaule
1999
S. ancophilum
1999-2000
8
S. bi/1-hookerii
2
1997-98-1999
S. cajamarquense
1999
S. cantense
1997-98
S. chiquidenum
2000
S. chomatophilum
8
1999-2000
S. gracilifrons
2000
S. hastiforme
9
1999-2000
S.huancabambense
2
1999-2000
S. hypacrarthrum
7
1999-2000
S. medians
5
2000
S. mochiquense
1999
S. orophylum
3
1999-2000
S. paucissectum
3
1999-2000
S. piurae
2
1999
S. raquialatum
1999
S. simplicissimum
3
1999
S. sogarandinum
10
1999-2000
S. wittmackii
Wild Solanum (Basarthrum)
(31 isolates)
2 14
1998 1-992-20003
13
S. caripense
1
1997
S. montanum
Wild tomatoes (31 isolates)
3
3
8
1998-99-2000
Lycopersicum hirsutum
6
1
5
1998-99
L. peruvianum
3
2000
L. pennelii
1
41
Host specificity
Pathogenic aggressiveness
Host specificity was tested by characterizing isolates of P. infestans from wild and
cultivated Solanaceae with molecular
markers: glucose-6-phosphate isomerase
(Gpi) and peptidase (Pep) allozymes,
restriction fragment length polymorphism
(RFLP) and amplified fragment length
polymorphism (AFLP) DNA fingerprinting,
and by assessing pathogenic fitness on
inoculated leaflets.
DNA fingerprinting
We obtained RFLP fingerprints for 239
isolates using the moderately repetitive
probe RG57 (Goodwin et al., 1992).
Hybridization and detection were conducted using the non-radioaC:tive kit ECL
(Amersham, lnc., Piscataway, NJ, USA)
according to the manufacturer's instructions.
Allozyme tests
A subset of n = 78 isolates (all of them
obtained in 1999) was analyzed for their
Gpi and Pep genotypes, using one of three
techniques: (1) cellulose acetate electrophoresis (CAE), (2) starch gel electrophoresis, or (3) polyacrylamide gel electrophoresis (PAGE). CAE was the simplest
technique for determining certain Gpi and
Pep genotypes (Goodwin et al., 1995), but
lacked resolution for differentiation of all
known genotypes. Starch gel electrophoresis for Gpi (Spielman et al., 1990) and PAGE
(Pep) were used when greater resolution was
needed (Vega-Snchez et al., 2000).
42
Research on Potato
Results
Characterization of isolates
markers
by molecular
Table 2. Number, location, and host origin of RG57 lineages defined by RFLP of isolates of Phytophthora
infestans collected in Peru between 1998 and 2000.
lsolates per host
Bred
Native
Wild
S. caripense
Wild
RG57 lineage1
Location 2
potatoes potatoes potatoes + Basarthrum tomatoes
EC-1
78
30
53
11
11
An, Ca,
(1110101001001101000111011)
Hca, Lib, Li,
Pi
PE-3
12
4
o
Ca, Li, Pi
(1100100001001100100111011)
US-1
2
14
Ca, Pi
o
o
4
(1010101011001101000110011)
PE-7
Ca, Hca, Li
o
3
o
10
3
(1110101001001100101111011)
Total per host
90
37
59
25
26
Ca, Li,
Ca, Hca,
Location per host
An, Ca,
Ca, Hca, An, Ca,
Hca, Lib, Lib, Li, Pi Hca, Lib,
Lib, Pi
Jun, Li,
Li, Pi
Li, Pi
Lib, Pi,
1
2 Usted by department code. An: Ancash, Ca: Cajamarca, Hca: Huancavelica, Jun: Junn, Lib: La Libertad, Li: Lima, and
Pi: Piura.
CIP Program Report 1999 - 2000
43
Discussion
Our results confirm and extend those of an
earlier study in which lineages EC-1, PE-3,
and US-1 were found among about 300
isolates coming from cultivated potatoes
in central and southern Peru (Prez et al.,
1999). Our sampling involved more host
species and we found another clonal
lineage, PE-7, principally associated with
wild tomatoes. The high degree of similarity in the results of the two studies supports
the hypothesis that our knowledge of the
Peruvian population of P. infestans is now
fairly accurate.
In general, our results are consistent with
the hypothesis that cultivated and wild
potatoes in Peru are attacked by the same
population of P. infestans. We found no
evidence to support host specificity among
wild and cultivated species within section
Petota of the genus Solanum. These res u lts
are also consistent with those of a recent
study of wild and cultivated potatoes in
Mexico. In that study, the same population
appeared to attack all host species that
were studied (Grnwald et al., 2000).
We did find evidence for host specificity
among cu ltivated potatoes and more
distant relatives within the genus Solanum.
The primary patato lineage of P. infestans
in Peru, EC-1, is not the primary lineage
attacking wild tomatoes or, at times,
S. caripense.
Table 3. Collection date, location, and host differentiation of isolates of P. infestans belonging to the EC-1
and US-1 lineages, collected from cultivated potatoes and S. caripense.
RG57 Lineage1
EC-1 (%)
US-1(%)
Groups
Year collected
Location
o
Bred potatoes
1999
Piura
50 (4)
Bred potatoes
2000
Piura
70 (9)
o
Bred potatoes
2000
Cajamarca
100 (10)
o
15
S. caripense
1998
Cajamarca
85 (6)
S. caripense
2000
Cajamarca
100 (5)
o
11
83 (1 O)
S. caripense
1999
Piura
100 (4)
S. caripense
2000
Piura
o
1 Number in parenthesis is the number of isolates per group.
44
Research on Potato
(cm)
3 ..-----------------~
A
2 ....
4 3 -
2 -
1 ....
1 -
P50.0001
p 50.0001
lsolates from S. caripense
- - lsolates from bred potatoes
2 -
p 50.0001
p 50.083
lsolates from S. huacabambense
- - lsolates from bred potatoes
-----
3
2
p 50.47
lsolates from S. sogarandinum
- - lsolates from bred potatoes
Figure 1. Lesion diameter on leaflets of wild Solanum and bred potatoes, 5 days after inoculation with a drop of
inoculum (5 x 103 sporangia/ml) of Phytophthora intestans from both hosts in detached-leaf inoculation assays.
(A), (B), (C): Solanum caripense vs bred potatoes. (D), (E), (F): wild potatoes vs bred potatoes. The isolates are
of EC-1 lineage except for (B) isolates of US-1 lineage from S. caripense, and (C) isolates of US-1 lineage from
both hosts. For each detached-leaf inoculation assay, the interaction is significant when Ps._ 0.05. Key: - =
bred potatoes, - = wild Solanum.
In sorne cases, host specificity between
cultivated potatoes and S. caripense was
associated with pathogen clonal lineage.
EC-1 was found on patato and US-1 on
S. caripense. This is similar to results
published for Ecuador (Erselius et al.,
1999), where EC-1 was also found on
patato and US-1 on S. caripense. Unlike in
Ecuador, however, EC-1 was also found on
S. caripense in our study. Nonetheless,
these EC-1 isolates appeared to be more
aggressive on S. caripense than on patato,
based on the significant interaction in
45
Conclusion
These results have implications for patato
breeding. Based on the results of this study,
it appears that breeders can use the
species we studied in section Petota as a
source of traits for introgression into
cultivated patato. There is little evidence
that a host specificity factor may be
confused with resistance to P. infestans. In
theory, patato breeders should be able to
use the potato popu lation of the pathogen
(the most abundant and easiest to use) to
screen a segregating population of potato
genotypes derived from a wi Id by cu ltivated cross. This has been the classical
approach to plant breeding. Thus, our
results support traditional breeding
techniques.
On the other hand, attempts to introgress
traits i nto cu ltivated potato from more
distant species in the genus Solanum could
cause problems. Our results and those of
others, (Erselius et al., 1999), indicate that
there are different populations of the
pathogen attacki ng these hosts in South
America. Breeders could potentially
introgress a host specificity factor into
cultivated potato, which would make it
appear resistant to the potato population of
the pathogen. Once planted in the proximity of the wi Id progenitor, however, the
putatively resistant patato could be
vulnerable to the pathogen population
specific to the wi Id progenitor.
46
Research on Potato
Acknowledgments
We want to thank Dr. William Fry of
Cornell University, USA, for providing the
probe RG57 and also several of the CIPLima staff. These are Dr. Bodo Trognitz for
providing the isolates of P. infestans of
1997-98 collection; Soledad Gamboa for
her assistance, as wel 1 as al 1 the techn ical
staff of the laboratory of P. infestans;
F. Mendiburu for statistical analyses; and
the staff in CIP-Huancayo for plant maintenance. This work was financially
supported by the French Ministry of
Foreign Affairs.
References
Deahl, K.L., L.R. Cooke, D.S. Shaw, and
D.J. Carlisle. 2000. Characterization of
isolates of Phytophthora infestans from
wild Solanum spp. in the UK.
Phytopathology 90(6):51 9.
Erselius, L.J., H.R. Hohl, M.E. Ordez,
P.J. Oyarzun, F. Jarrin, A. Velasco,
M.P. Ramn, and G.A. Forbes. 1999.
Genetic diversity among isolates of
Phytophthora infestans from various
hosts in Ecuador. In: lmpact on a
changing world: Program report, 199798. lnternational Patato Center Lima
Peru. p. 39-48.
'
'
47
49
Research on Potato
Table 1. Solanum accessions evaluated far resistance to late blight, Lima, Peru, 1999-2000.
Series and s~ecies
2n EBN 1 Country
No.
m
s
o Proportion No.
Collectors
CIP No.
of origin tested 2
resistant selected
No.
genotypes
(m+r+O)
/No.
ACAULIA
S. a/bicans (alb)
CIP 761452 OCH 12089
72
4 PERU
CIP 762120 OCH 14789
72
4 PERU
CIP 762584 OCHS 16028
72
4 PERU
CIRCAEIFOLIA
S.circaeffolium(crc)
CIP 761344 OCHS 11909
24
1 BOLIVIA
S. circaeifolium var. capsicibaccatum (cap)
CIP 761030 OCHS 11915
24
1 BOLIVIA
CIP 761030 OCHS 11915
24
1 BOLIVIA
CIP 762303 OCHS 15489
24
1 BOLIVIA
COMMERSONIANA
S. commersonii (cmm)
CIP 761087 FB 401 O
URUGUAY
24
CIP 762454 URY 4
URUGUAY
24
CIP 762475 URY 31
24
URUGUAY
CON ICIBACCATA
S.chomatophilum(chm)
CIP 761587 OCH 13341
24
2 PERU
CIP 762568 OCHS 12553
24
2 PERU
CIP 762611 OCHS 16072
24
2 PERU
S. colombianum (col)
CIP 762793 SCL 5050
48
2 ECUADOR
CIP 762793 SCL 5050
48
2 ECUADOR
S. flahaultii (flh)
CIP 761877 OCH 14105
COLOMBIA
48
CIP 761878 OCH 14106
48
COLOMBIA
S. irosinum (irs)
CIP 761252 OCH 11640
24
2 PERU
CIP 762257 OCHS 15210
24
2 PERU
S. paucijugum (pcj)
ECUADOR
CIP 762800 SCL 5084
48
ECUADOR
CIP 762803 SCL 5096b
48
48
ECUADOR
CIP 762816 SCLG 5151
S. urubambae (uru)
CIP 761044 OCH 13781
24
2 PERU
CIP 762368 OCHS 15654
24
2 PERU
S. violaceimarmoratum (vio)
CIP 760563 VSOA 7
24
2 BOLIVIA
DEMISSA
S. demissum (dms)
CIP 761893 OCH 14154
72
4 MEXICO
CIP 761050 OCH 14218
72
4 MEXICO
S. hougasii (hou)
CIP 761902 OCH 14172
4 MEXICO
72
CIP 761899 OCH 14168
4 MEXICO
72
o o
o o
o o o
0.21
0.02
O.DO
o
o
o
0.18
27
31
0.98
1.00
0.77
42
11
24
48
4
48
48
48
42
19
5
4
20
9
2
7
17
0.13
0.60
0.90
3
19
23
48
48
48
47
35
46
1
12
0.02
0.27
0.04
o
o
o
48
48
10
43
36
5
2
o
o o
0.79
0.10
48
47
11
24
31
20
4
2
2
1
0.77
0.49
o
o
o
4
48
48
40
18
2
2
6
28
o
o
0.17
0.63
o
o
42
39
37
26
7
21
14
14
16
o
o
o o
0.38
0.76
0.43
o
o
o
48
48
18
48
48
48
38
47
48
10
1
51
42
56
48
48
48
44
48
48
48
o
o
o
o
o o
2
17
o o
1 o
1
2
8
o
1
5
10
43
1.00
0.63
20
15
22
10
0.98
1 26
13 35
17
1.00
1.00
38
28
36
37
1.00
1.00
17
26
o
o
12
11
51
Table 1. (continued)
Series and s~ecies
Collectors
CIP No.
No.
2n
No.
EBN 1 Country
of origin tested2
Proportion
No.
resistant selected 3
genotypes
(m+r+O)
/No.
S. iopetalum (iop)
72
CIP 761928 OCH 14208
4 MEXICO
72
CIP 761935 OCH 14221
4 MEXICO
72
CIP 761930 OCH 14212
4 MEXICO
LIGNICAULIA
S. lignicaule (lgl)
24
CIP 761210 OCH 11315
PERU
CIP 761236 OCH 11617
24
PERU
CIP 761652 OCH 13585
24
PERU
LONGIPEDICELLATA
S. fendlerii (len)
48
CIP 761921 OCH 14199
2 MEXICO
CIP 761923 OCH 14202
48
2 MEXICO
48
CIP 761926 OCH 14205
2 MEXICO
S. stoloniferum (sto)
CIP 761884 OCH 14135
48
2 MEXICO
CIP 761062 OCH 14145
48
2 MEXICO
MEGISTACROLOBA
S. do/ichocremastrum (dcm)
CIP 761043 OCH 12071
24
1 PERU
CIP 761439 OCH 12074
24
1 PERU
CIP 761470 OCH 13013
24
1 PERU
CIP 761470 OCH 13013
24
1 PERU
S. megistacrolobum subsp. toralapanum (mga)
24
CIP 760535 HAM 200
2 BOLIVIA
24
CIP 761109 OCH 7609
2 PERU
CIP 761403 OCH 12032
24
2 BOLIVIA
S. megistacrolobum subsp. toralapanum (tor)
CIP 760459 HHA 6616
24
2 BOLIVIA
24
2 BOLIVIA
CIP 761369 OCH 11964
24
2 BOLIVIA
CIP 761055 OCHS 11914
S. raphanifolium (rap)
24
2 PERU
CIP 761113 OCH 7613
24
CIP 761640 OCH 13572
2 PERU
24
CIP 761671 OCH 13610
2 PERU
S. sogarandinum (sgr)
24
CIP 761586 OCH 13336
2 PERU
CIP 762410 OCHS 15723
24
2 PERU
PINNATISECTA
S. cardiophy/Jum (cph)
CIP 762561 OCH 14157
24
MEXICO
PIURANA
S. cantense (cnt)
CIP 762241 OCHS 15159
24
2 PERU
S. chiquidenum (chq)
24
CIP 761588 OCH 13345
2 PERU
CIP 761870 OCH 13963
24
2 PERU
CIP 762573 OCHS 12566
24
2 PERU
52
Research on Potato
o
o
o o
o o
48
48
48
11
21
16
48
47
48
29
12
21
17
27
24
48
45
48
9
45
48
44
48
48
48
1.00
1.00
0.77
23
46
24
2
8
3
o
o
o
0.40
0.74
0.56
o
o
o
31
o o o
o o o
13
11
o o
0.81
34
O.DO
O.DO
o
o
17
48
0.93
1.00
3
30
O.DO
o
o
48
47
48
48
48
47
45
48
43
48
48
1
27
47
o o o
o o o
o
1 2
o o o
o 21 21
o
15
6
1 o o
48
27
32
1
25
10
5 16 26
1
1 o
o
16
6
0.98
0.07
0.69
o
o
48
48
48
1
3
5
26
29
29
17
14
9
4
2
5
0.98
0.94
0.90
15
24
17
48
47
1
42
10
4
10
27
1
0.98
0.11
16
2
29
o o
29
1.00
12
48
22
1.00
48
48
48
37
33
35
11
15
11
o o
o o
o
2
0.23
0.31
0.27
.12
l1
14
21
0.00
0.06
0.00
0.98
0.44
0.02
o
16
o
8
1
Table 1. (continued)
Series and s~ecies
CIP No.
Collector's
No.
2n
Proportion
No.
resistant selected 3
genotypes
(m+r+O)
/NO.
S. humectophilum (hmp)
CIP 761052 OCH 11753
S. hypacrarthrum (her)
CIP 761259 OCH 11692
CIP 761204 OCHS 11308
CIP 762104 OCHS 14715
S. paucissectum (psc)
CIP 761247 OCH 11634
CIP 762124 OCHS 14816
S. piurae (pur)
CIP 761868 OCH 13959
CIP 761072 OCHS 11615
POLIADENIA
S. polyadenium (pld)
CIP 761911 OCH 14187
TUBEROSA
S. alandiae (aln)
CIP 760469 HHA 6657
CIP 760473 HHA 6665
CIP 761394 OCH 12013
S. ambosinum (amb)
CIP 761842 OCH 13852
CIP 761194 OCHS 11298
CIP 761053 OCHS 11865
S. brevicaule (brc)
CIP 760461 HHA 6619
CIP 760479 HHA 6690
CIP 761074 OCH 11934
S. bukasovii (buk)
CIP 761120 OCH 7717
CIP 761517 OCH 13166
CIP 761664 OCH 13602
CIP 761720 OCH 13679a
CIP 761805 OCH 13796
CIP 761806 OCH 13798
CIP 761847 OCH 13858
CIP 762434 OCH 15822
CIP 761152 OCHS 10114
CIP 761301 OCHS 11851
S. cajamarquense (cjm)
CIP 762616 OCHS 16118
CIP 762619 OCHS 16121
S. candolleanum (cnd)
CIP 761041 OCHS 11913
CIP 762168 OCHS 14959
CIP 762185 OCHS 15011
24
PERU
53
24
24
24
PERU
PERU
PERU
48
23
48
24
24
2 PERU
2 PERU
44
47
24
24
2 PERU
2 PERU
48
48
o
11
o
15
2
o o o
23
41
6
1 o
41
2
1 o
26
o 13
8
o o 2 46
o o o 48
50
24
MEXICO
48
24
24
24
BOLIVIA
BOLIVIA
BOLIVIA
48
48
48
28
48
48
24
24
24
2 PERU
2 PERU
2 PERU
48
48
48
48
47
48
24
24
24
2 BOLIVIA
2 BOLIVIA
2 BOLIVIA
48
48
48
48
47
48
24
24
24
24
24
24
24
24
24
24
2
2
2
2
2
2
2
2
2
2
PERU
PERU
PERU
PERU
PERU
PERU
PERU
PERU
PERU
PERU
48
48
48
48
48
47
48
30
48
43
30
20
47
35
47
26
33
30
47
10
24
24
PERU
PERU
48
48
1
2
24
24
24
2 BOLIVIA
2 BOLIVIA
2 BOLIVIA
48
48
48
48
48
48
o o
48
o o
o o o
o o o
o o o
1
o o
o o o
20
0.06
0.46
O.DO
0.15
0.07
0.83
o
o
o
o
o
1.00
1.00
20
4
1.00
12
0.42
O.DO
O.DO
18
o
o
O.DO
0.02
O.DO
o
o
o o o
1
o o
o o o
17
1 o
o
23
5
1
o o
o
10
3
o 1 o
17
o
4
13
o
2
o o o
1
o o
26
o
7
O.DO
0.02
O.DO
o
o
o
0.38
0.58
0.02
0.27
0.02
0.45
0.31
O.DO
0.02
0.77
7
1
7
2
0.98
0.96
18
11
o o o
o o o
o o o
O.DO
O.DO
O.DO
10
14
30
30
o
3
o
o
2
o
o
o
3
8
53
Table 1. (continued)
Series and s~ecies
CIP No.
Collector's
No.
2n
S. coelestipetalum (cap)
24
CIP 761660 OCH 13596
24
CIP 761728 OCH 13686
24
CIP 762005 OCH 14351
S. huancabambense (hcb)
24
CIP 761238 OCH 11619
24
CIP 761238 OCH 11619
24
CIP 761239 OCH 11626
24
CIP762123 OCHS 14815
S. huarochiriense (hro)
24
CIP 761215 OCH 11325
CIP 761224 OCH 11335
24
24
CIP 761224 OCH 11335
24
CIP 761265 OCH 11699
S. leptophyes (lph)
24
CIP 761766 OCH 13730
24
CIP 760805 VSAL 126
24
CIP 760895 VSH 248
S. marinasense (mrn)
24
CIP 761669 OCH 13608
24
CIP 761774 OCH 13737
24
CIP 761812 OCH 13809
S. medians var. autumnale (aut)
24
CIP 761198 OCHS 11302
CIP 762629 OCHS 12573
24
24
CIP 762256 OCHS 15205
S. microdontum (mcd)
24
CIP 762314 OCHS 15534
24
CIP 760531 HAM 176
CIP 760534 HAM 187
24
S. mochiquense (mcq)
CIP 761037 OCHS 14870
24
S. multiinterruptum (mtp)
CIP 761260 OCH 11693
24
24
CIP 761422 OCHS 12055
24
CIP 762405 OCHS 15716
S. oplocense (opl)
CIP 761352 OCH 11927
24
48
CIP 761362 OCH 11947
48
CIP 761362 OCH 11947
S. orophilum (orp)
24
CIP 761440 OCH 12077
CIP 761443 OCH 12080
24
CIP 761475 OCH 13020
24
S. sparsipilum (spl)
CIP 760475 HHA 6669a
24
54
Research on Patato
EBN 1
No.
Country
of origin tested2
Proportion
No.
resistant selected 3
genotypes
(m+r+O)
/No.
o
6
3
o o o
o 1 o
0.19
0.00
0.02
o
o
o
o
o
o
o
o
o
o
o
0.35
0.15
0.46
0.06
9
5
20
o o
1 o
1 o
0.18
0.13
0.44
6
39
42
33
9
5
0.88
0.19
0.13
48
48
48
37
45
16
6
1
17
5
2
15
PERU
48
11
25
12
2 PERU
2 PERU
2 PERU
48
48
48
26
2
19
18
9
27
2
24
2
2 BOLIVIA
4 BOLIVIA
4 BOLIVIA
45
44
48
11
44
48
2 PERU
2 PERU
2 PERU
48
48
48
o
1
2
17
37
22
2 BOLIVIA
48
46
2 PERU
2 PERU
2 PERU
48
48
48
39
48
47
2
2
2
2
PERU
PERU
PERU
PERU
48
48
48
48
31
41
26
45
2
2
2
2
PERU
PERU
PERU
PERU
34
48
48
48
33
44
46
48
2 PERU
2 BOLIVIA
2 BOLIVIA
48
46
46
48
41
38
2 PERU
2 PERU
2 PERU
51
47
48
42
41
27
2 PERU
2 PERU
2 PERU
48
48
48
2 BOLIVIA
2 BOLIVIA
2 BOLIVIA
14
4
10
2
3
3
12
1
o
o
o
o o
o o
o
5
o
8
1
4
2
o o
1 o
6
3
o
5
o
o
o
1
3
o
o
o
o
o
2
0.23
0.06
0.67
15
6
38
0.77
2
13
0.46
0.96
0.60
3
14
o
o
24
o o
10
0.00
0.11
0.17
o
o
o
o
o
o
26
8
o o o
o o o
28
0.03
0.08
0.04
0.00
o
o
0.76
0.00
0.00
o
8
o
o
1.00
0.98
0.96
1
2
0.04
Table 1. (continued)
Series and s~ecies
CIP No.
Collector's
No.
2n
EBN 1 Country
No.
of origin tested 2
Proportion
No.
resistant selected 3
genotypes
(m+r+O)
/No.
S. velardei (vlr)
CIP 761730 OCH 13688
CIP 762027 OCH 14387
CIP 762028 OCH 14387a
S. wittmackii (wtm)
CIP 761566 OCH 13267
CIP 761205 OCHS 11309
CIP 762077 OCHS 14626
YUNGASENSA
S. berthaultii (ber)
CIP 761390 OCH 12008
S. chacoense (che)
CIP 761399 OCH 12026
CIP 762270 OCH 15266b
CIP 762274 OCH 15271
S. tarijense (lar)
CIP 761007 OCH 12001
CIP 762334 OCHS 15596
24
PERU
PERU
PERU
48
48
48
20
23
32
23
25
15
o
5
o o
1 o
0.58
0.52
0.33
24
24
24
PERU
PERU
PERU
48
48
48
12
26
47
32
21
1
3
1
o
o o
0.75
0.46
0.02
24
2 BOLIVIA
44
23
15
0.48
15
24
24
24
2 BOLIVIA
2 PARAGUAY
2 PARAGUAY
44
48
48
27
26
41
10
16
7
7
5
0.39
0.46
0.15
5
3
24
24
2 BOLIVIA
2 BOLIVIA
Total
24
48
48
6306
o o
o
4
o o
9
35
47
1
3532 1316 685 743
0.81
0.02
o
o
o
o
o
o
10
3
801
55
r + O , m + r , and _Q , respectively,
n
n
n
m +
Results
lnitial symptoms of infection were observed three days after inoculation, and
susceptible individuals reached 100%
56
Research on Potato
f:
**
11
O>
e 10
::.i2
9
e
e:
(1)
.f
c..
X
(1)
(1)
(/)
(1)
:::J
i=
4
3
2
* *
(1)
"O
11
..
7
6
13
12
10 L
O>
e
9 ::.i2
8 e
7
6
5
4
3
2
1
e:
.fE
(1)
c..
X
(1)
(j
.o
:::J
1-
o
o
2 3 4 5 6 7 8 9 10 11 12 13
Field experiment ranking
57
"*-e
0.9
:e
0.8
::::J
:;:
-~
e:ro
()
"C
.....
~.._
* *
...
'
0.7 -
"-l ,,
0.6 ,_
>
.s
0.5 ,_
0.4
0.3
~
ro
~o.
a..
0.2
0.1
~~
,.,,,
**
-~
***
__
,,
~-
.,
MJo:
~*
~~
1
li
-:;:
:e
(])
- 0.6
:e
:g_
0.5
o
en
::::J
en
- 0.4
0.3
:eo
- 0.2
a..
(])
e:
._,
*'
f!"I***
..
en
(ij
"O
- 0.7
>
; * * * "'
,.**
0.9
- 0.8
*-
::::J
'*
"'
** "'
.,,,
Q)
~
* '*
..
*
*
1.0
'*
)~.
11
I!
11
o.
- 0.1
1
1
1
O.O
F_igure 2. P!o~ortion~ of individuals in each o'.139 germplasm samples tested in true-seed experiments falling
mt~ quant1tat1ve res1st~nce (m + r), susceptible (s) and qualitative resistance (O) categories. (* = quantitatively
res1stant; = susceptible; 1 = qualitatively resistant).
S.
S.
S.
S.
S.
S.
S.
58
Research on Potato
confirming the likely presence of quantitative resistance. Eight hundred and one
promising genotypes in 35 species of 11
taxonomic series were selected by comparison of the results obtained from the
inoculation of plants grown from true
seed and from tubers. These selections
are currently being propagated for resistance evaluation at the genotypic level
(Table 1 ).
Discussion
Limitations to the greenhouse assay
The greenhouse assay was a convenient
way to evaluate large samples of
germplasm under applied disease pressure.
However, susceptible individuals reached
100% foliar infection only eight days after
inoculation. In contrast, it takes 3-4 weeks
for moderately resistant or susceptible
clones to reach 80-100% infection in the
field. The rapid greenhouse epidemics
may not provide the best conditions to
59
60
Research on Potato
S.
S.
S.
S.
urubambae/ S. violaceimarmoratum/
cantense/ S. cajamarquense,
orophilum, S. velardei and
wittmackii. We therefore report the first
S.
S.
S.
S.
S.
violaceimarmoratum,
cajamarquense, S. coelestispetalum,
medians, S. microdontum,
multiinterruptum, S. oplocense, and
orophilum). This may facilitate their use
in breeding.
This population approach to germplasm
evaluation provides important infarmation
needed to select and optimize strategies
for the use of new resistance sources in
breeding. For example this effort will
orient the selection of new donors of
resistance for tests of heritability and
complementarity with currently deployed
resistance types. The use of multiple
genotypes of a given donor species and a
carefully chosen set of diverse recurrent
parents may help circumvent the poorly
understood genetic interactions between
wild and cultivated species that often
result in fertility problems in F1 and F2
generations (Santini et al., 2000). The
selection of diverse donar species using
taxonomic, ecogeographic and mechanistic criteria should also contribute to
building complex resistance types that are
not readily overcome by the pathogen.
Acknowledgments
This work was supported in part by the
Department for lnternational Development
(DFID), United Kingdom. We thank E. de
la Torre, H. Ponce, G. Marticorena and
O. Gaspar far technical assistance in the
greenhouse and laboratory, V. Otazu far
input on design and execution of the
experiments, C. Arellano for statistical
consultation, and R. Hijmans, A. Panta,
and E. Mihovilovich for helpful comments
on the manuscript.
References
Colon, L.T. and Budding, D.J. 1988.
Resistance to late bl ight (Phytophthora
infestans) in ten wild Solanum species.
Euphytica Supplement p. 77-86.
Colon, L.T., D.J. Budding, L.C.P. Keizer,
and M.J.J. Pieters. 1995. Components of
resistance to late blight (Phytophthora
infestans) in eight South American
Solanum species. European Journal of
Plant Pathology 101 :441-456.
Ewing E.E., l. Simko, C.D. Smart,
M.W. Bonierbale, E.S.G. Mizubuti,
G.D. May, and W.E. Fry. 2000. Genetic
mapping of qualitative and quantitative
field resistance to Phytophthora
infestans in a population derived from
Solanum tuberosum and Solanum
berthaultii. Molecular Breeding
6:25-36.
Fry, W.E. 1978. Quantification of
general resistan ce of patato cu ltivars
and fungicide effects for integrated
control of late blight. Phytopathology
68:1650-1655
61
62
Research on Potato
Monitoring the genetic variation for horizontal resistance to late blight in the
most advanced sources of horizontal resistance to late blight-population B
group three cycle one (B3C1 )-has been carried out at CI P by assessing the
amount of genetic variance for resistance and tuber yield. A random sample
of resistant clones from B3C1 were used to obtain progenies according to
mating designs NC Design 1 and Line x Tester. Resulting progenies were
evaluated for resistance to late blight and total tuber yield in the field under
high disease pressure at Comas, Peru. Three independent estimates of heritability, including parent-offspring regression for area under the disease
progress curve (taken as :.. parameter for resistance), were high and reasonably close (h 2 = 0.48, h2 = 0.53, and h2 = 0.40). Additive genetic variance and
heritability for resistance in this population are large enough to ensure further
progress from selection.
63
64
Research on Potato
Results
c.v.= 9.8%
df
1
2
15
17
35
MS
87714.694
264299.194
71709.428
16088.694
5.45 **
3.68 **
4.46 **
65
Table 4. Components of additive genetic variances and heritability estimates far AUDPC and tuber yield in
samples of B3C1 late blight resistant clones.
AUDPC
Tuber yield
h2
h2
Mating design
SE
SE
100384
0.48
0.45
3.78
0.10
0.17
Design 1
0.18
0.25
0.08
281082
0.53
65.40
Line x tester
Parent-offspring
19407166
0.40
0.06
Nurnber
50
45
Mean: 2556
sd: 877
40
35
30
25
20
15
10
0500
5011000
10011500
1501-2000
2001-2500
2501-3000
AUDPC
66
Research on Patato
3001-3500
3501-4000
References
Forbes, G.A. 1998. Genotype by
environment reaction of potato to the
late blight pathogen. In: lmpact on a
67
68
Research on Potato
846 p.
Turkensteen, L.J. 1993. Durable resistance
of potatoes against Phytophthora
infestans. In: Jacobs, T. and J.E.
Parlevliet (eds.). Durability of disease
resistance. Kluwer Academic Publishers,
Dordrecht, Netherlands. p. 11 5-1 24.
Van der Plank, J.E. 1968. Disease
resistance in plants. Academic Press,
NY, USA. 206 p.
Waistie, R.L. 1991. Breeding for
resistance. In: lngram, D.S. and P.H.
Wil 1iams (eds.). Phytophthora infestans:
The cause of late bl ight of pota to.
Academic Press, San Diego, CA, USA.
p. 193-224.
69
70
Research on Potato
Disease severity
Disease severity was estimated visually
every 4 or S days and expressed as the
fraction of the foliage infected. Relative
area under the disease progress curve
(rAUDPC) was calculated according to
Fry (1978).
Results
Disease
We observed LB at both sites, but rAU DPC
was much higher in Oxapampa where the
conditions were more conducive to
disease development (Table 1). The effects
on disease of variety, N rate, F treatment,
and the interactions of variety*N,
variety*F, and N*F were al 1 significant at
P = O.OS at both sites. In Oxapampa there
were no treatments without LB, whereas in
Huancayo there was LB only in the
treatments without fungicides (Figures 1
and 2). Because of space limitations, only
the results far the lowest (null) and highest
(4- or 7-day interval) fungicide treatments
are shown (Figures 1 and 2).
Late blight severity increased with N but
the effect depended on disease pressure,
varietal resistance, and fungicide use.
There was no effect of N on disease at
very high or low levels of disease (Figures
1A, 1 G, 2A, 2G). At intermediate levels of
disease, LB severity was lower at NO than
at Nl 60 and N320 far Yungay (Figures 2C
and 2E). For Amarilis, however, there were
significant differences between NO and
N320, but Nl 60 could not be statistically
distinguished from NO or N320 (Figures 1 C
and 1 E).
71
Table 1. Average rAUDPC1, IR2, and yield 3 using various treatments 4 in varieties Amarilis
Oxapampa and Huancayo, Peru, 1999/2000 growing season.
NO
N160
f"
F10
FF10
F4
FOxapampa
f4
Amarilis
0.6
0.6
rAUDPC
0.1
0.7
0.1
0.7
0.7
83
372
37
96
487
37
IR
34
o.o . 10.5
o.o
O.O
O.O
Tuber yield
o.o
18.6
Yungay
0.6
0.8
0.6
0.8
rAUDPC
0.7
0.1
0.2
87
83
420
32
22
IR
26
496
o.o 11.8
O.O
O.O
O.O
Tuber yield
O.O
11.9
Huancayo
FAmarilis
rAUDPC
O.O
IR
193
Tuber yield
3.8
Yungay
rAUDPC
0.1
IR
175
Tuber yield
2.9
f 14
f7
f-
F14
F7
F-
and Yungay in
N320
F10
0.6
79
O.O
F4
0.2
511
17.4
0.6
84
0.3
504
11.5
F14
f7
o.o
O.O
O.O
O.O
O.O
O.O
0.1
O.O
O.O
232
5.6
242
5.6
345
8.9
379
8.9
391
11.1
321
6.5
319
5.9
407
11.7
O.O
O.O
O.O
436
9.0
0.2
275
2.7
O.O
401
9.3
426
8.3
447
11.8
O.O
O.O
191
4.5
224
5.1
0.3
277
3.1
Treatment: NO = zero nitrogen (N) fertilization; N160 = 160 kg/ha; N320 = 320 kg/ha; F- = no fungicide;
F4 = fungicide every 4 d; F7 = fungicide every 7 d; F10 = fungicide applied every 10 days; F14 =
fungicide every 14 days.
72
Research on Potato
Blight proportion
1.00
Ground cover
0.80
0.60
0.60
0 ..40
0.40
O.al
02)
0.00
0.00
20
60
40
80
1.00
Oxaparnpe
F4
0.80
120
100
1.00
0.60
0.60
0.40
0.20
0.20
000
000
20
60
40
80
100
40
60
100
80
,---:-;~-=:::;:::;;;;::::::::-:----
120
1.00 . , . - - - - - - - - - - - - - - - - - - - - .
Huancayo
f.
0.80
20
O.BO
0.40
20
40
60
80
100
120
1.00 - . - - - - - - - - - - - - - - - - - - - - - - .
0.8J
0.60
0.60
0.40
0.40
0.20
020
0.00
1.00 . . . . - - - - - - - - - - - - - - - - - - - .
OUpampa
0.8J
f.
Oxapampa
.J-..................................~~;:~;as;~S:;S;;~::J
20
40
60
80
100
120
0.00
140
25
00
75
100
125
150
25
50
75
100
125
150
G
1.00
1.00
Huancayo
F7
0.80
0.80
0.60
0.60
0.40
0.40
a aa
0.20
0.20
000
000
20
40
60
80
100
120
140
o NO
N160
N320
Figure 1. Blight proportion and ground cover for Amarilis, three levels of nitrogen (N) fertilization and different
levels of fungicides, in Oxapampa and Huancayo. Vertical bars indicate the standard error of the mean. Key: NO
= zero nitrogen (N) fertilization, N160 = 160 kg N/ha, N320 = 320 kg N/ha, F- = no fungicide, F4 =
fungicide applied every 4 d, F7 = fungicide every 7 d.
N320 there was a strong yield loss due to
LB for Amarilis. lt is not clear whether the
differences in the response of Yungay and
Amarilis are related to the observed level
of infection (a function of their resistance
73
Blight proportion
1.0
1.0
0.8
0.8
0.6
0.6
0.4
0.4
0.2
0.2
o.o
O.O
20
40
60
1.0
D
0.8
a
a
0.6
0.4
0.2
0.2
o.o + - - s . - - - . - - - - , - - - - - r - - - . - - - - 1
20
80
60
40
1.0
--~--~-~-~--~---<
20
40
60
80
100
120
1.0
Hu ancayo
o.a
O.O
120
100
120
100
1.0
0.8
F-
60
0.6
0.4
Oxapampa
120
100
80
Ground cover
o.a
o.6
0.6
0.4
0.4
0.2
0.2
o.o
O.O
20
40
60
80
100
120
140
25
50
75
100
125
150
25
50
75
100
125
150
G
1.0
1.0
Huancayo
F1
0.8
0.8
0.6
0.6
0.4
0.4
a a a
0.2
o.o
0.2
+----..................~.............~....~:e:S~:S:S:~!l::ll
o
20
40
60
80
100
120
o.o
140
N320
Figure 2. Blight proportion and ground cover for Yungay, three levels of nitrogen (N) fertilization and different
levels of fungicides, in Oxapampa and Huancayo. Vertical bars indicate the standard error of the mean.
Treatment: NO = zero nitrogen (N) fertilization, N160 = 160 kg N/ha, N320 = 320 kg N/ha, F- = no fungicide,
F4 = fungicide applied every 4 d, F7 = fungicide every 7 d.
too low far Amarilis at N320/F-. This
apparent decrease in radiation use efficiency conflicts with results of Haverkort
and Bicamumpaka (1986) and Van Oijen
(1990).
74
Research on Potato
12
A(Amarilis)
10
12
N320 F7
10
N320 F-
(F7)
(F7)
4
(F-)
2
o
100
200
300
400
500
100
200
2
300
400
500
Figure 3. lntercepted radiation vs yield in Huancayo far Amarilis (A) (slope = 0.036) and Yungay (B) (slope =
0.026) at three levels of nitrogen (N) and two levels of fungicides. Oblique line is the regression between
intercepted radiation and yield (separate far each variety) far all observations in Huancayo. Arrows indicate the
effect of Non yield between levels of Nin the presence of late blight(F-), and absence of late blight(F7). Key: NO
= zero Nfertilization, N160 = 160 kg N/ha, N320 = 320 kg N/ha, F- = no fungicide, F7 = fungicide applied
every 7 d.
variety choice when aiming at increased
yields. Resistant varieties are always
useful where there is LB, but in sorne
cases, when farmers shift from low to high
levels of N fertilization, they might
become even more valuable. Possibly
there cou Id be a stronger effect of N on LB
at high N levels if P and K are low.
However, this scenario is not relevant
because low P and K would greatly reduce
yield, hence reducing these inputs is not a
viable management option.
For Yungay in Huancayo, the investment
in N fertilizer would only have been useful
if combined with fungicide use. This did
not hold for Amarilis, which has higher LB
resistance, and where LB only influenced
the effect of N on yield when an overdose
of N was supplied. For varieties such as
Amarilis, N does not seem to be an
important factor for integrated management of LB, because there is no evidence
of a need to adjust optimum N fertilization
because of the disease.
Acknowledgements
We thank Greg Forbes, Rebecca Nelson,
and Consuelo Arellano for reviewing this
paper.
References
Awan, A.B. and R.A. Struchtemeyer. 1957.
The effect of ferti 1ization on the
susceptibility of potatoes to late blight.
American Potato Journal 34:315-319.
Carnegie, S.F. and J. Colhoun. 1983.
Effects of plant nutrition on
susceptibility of potato leaves to
Phytophthora infestans.
Phytopathologische Zeitschrift
1 08:242-250.
Cohen, Y. and J. Rotem. 1987. Sporulation
of foliar pathogens. In: Pegg, G.F. and
P.G. Ayres (eds.), Fungal infection of
plants. Cambridge University Press, UK.
p. 314-333.
Crosier, W. 1934. Studies in the biology of
Phytophthora infestans (Mont.) de Bary.
Memoir 155. Cornell University
75
76
Research on Potato
Phytophthora infestans
77
78
Research on Patato
Table 1. Management practices used at each of the sites in evaluating late-blight development and patato
growth parameters in 1999 and 2000.
Year Management
Experimental site
1999 practice
Loreto, Short rain
Kabete, Long rain
Kalengyere, Season "B"
Planting date
29/10/1999
26/3/1999
16/10/1999
Fertilizer application
Diammonium phosphate Diammonium phosphate Diammonium phosphate
1nsecticide
Metasystox
Metasystox
Dimetheote
Fungicide
Dithane M-45
Dithane M-45
Dithane M-45
13(7/1999
Harvesting
02/2/1999
11/1/2000
Loreto, Long rain
Kabete, Long rain
2000
Kalengyere., Season "A"
Planting date
13/4/2000
12/4/2000
20/3/2000
Fertilizer application
Diammonium phosphate Diammonium phosphate NPK
Metasystox
Dimetheote
lnsecticide
Metasystox
Dithane M-45
Fungicide
Dithane M-45
Dithane M-45
Harvesting
26/7/2000
26/7/2000
23/7/2000
using an assessment scale of 0-100%. At
least five disease assessments were
recorded. At crop maturity, the incidence
of tuber blight was also quantified visually.
Tubers that did not show visual symptoms
were stored for three weeks and subsequently observed or plated to look for
additional incidence of tuber blight.
At the three experimental sites (Kabete,
Loreto, and Kalengyere), weather equipment (Hobo Pro Series, MA, USA, and
Watchdog Data Logger, Spectrum Technologies, Plainfield, IL, USA) monitored
environmental parameters such as temperature, relative humidity, rainfall, and
hours of sunshine or photosynthetic active
radiation. Additional data from the University of Nairobi weather station were also
used.
Data analysis
Mean values of disease incidence and
severity were calculated using SAS (1989).
Similarly, area under disease progress
curves (AUDPC) was calculated from
values for disease severity, as described by
Campbel 1 and Madden (1990). The
development and progress of late blight in
fungicide-treated plots versus untreated
controls were graphically compared using
AUDPC. The effects of the intervals
between fungicide applications on lateblight development were computed and
Results
Dynamics of late blight in relation to
fungicide application
During the 1999 cropping season, disease
severity at Kabete was low compared to
Loreto (Figure 1) The final disease level
(AU DPC) of AsanteNictoria was 250
(% disease days) at Kabete and 1200 at
Loreto. During the 2000 cropping season,
late-blight severity was very low at Loreto,
especial ly in fungicide-treated plots. In the
control (untreated) plots, the disease was
highest in the susceptible variety Kerr's
Pink (with an AU DPC val u e of 501 .1 ),
compared to Tigoni (AUDPC 163.9) and
AsanteNictoria (AUDPC 201.6) (Table 2).
79
Table 2. Effect of fungicide (Dithane M-45) application intervals on disease severity and patato yields of
patato at Loreto, long rains, and Kalengyere (season A, year 2000).
AUDPC2
Yield (Vha)
Tubers (no.)
Application intervals Variety1
Loreto, Kenya
Tigoni
0.2
57.0
495
7 days
0.4
349
59.8
7 days
AsanteNictoria
0.7
42.0
343
7 days
K. Pink
Tigoni
0.4
54.0
462
14 days
0.7
57.4
327
14 days
AsanteNictoria
0.9
14 days
K. Pink
41.5
223
Tigoni
21 days
3.6
52.5
400
21 days
5.2
56.0
254
AsanteNictoria
K. Pink
10.1
21 days
40.3
195
Tigoni
Control
163.9
51.2
329
201.6
Control
AsanteNictoria
53.9
199
K. Pink
501.1
35.2
17
Control
2.5
Mean
50.5
339
2.9
LSD 0.05
4.5
65.3
cv (%)
64.06
9.33
28.58
Kalengyere, Uganda
7 days
Rutuku
O.O
21.4
221.3
7 days
Ka bale
23.5
19.5
144.7
7 days
Victoria
19.6
22.5
200.3
14 days
Rutuku
2.1
23.2
214.7
14 days
Ka bale
52.5
19.5
176.0
14 days
Victoria
60.6
22.5
200.7
21 days
Rutuku
2.5
23.1
233.3
21 days
Kabale
69.3
19.5
172.3
21 days
Victoria
101.9
23.3
241.7
Tigoni
Control
90.3
21.0
234.0
Control
AsanteNictoria
185.9
17.3
164.3
Control
K. Pink
225.8
19.2
216.3
Mean
23.2
21.6
201.6
LSD 0.05
35.0
6.1
53.5
cv (%)
176.3
35.2
30.9
Note: Dithale M-45 (malcozeb) was applied at the rate of 3 kg/ha. A total of five applicatiols were made durilg the
croppilg seasol. Experimelts at Loreto were plalted Ol April 13 ald harvested Ol July 26. At Kalelgyere, plaltilg
was Ol March 20 ald harvestilg was July 23.
1 varieties Rutuku ald Tigoli are moderately resistalt ald Kabale is moderately susceptible, while AsalteMctoria ald Kerr
Pilk are susceptible to late blight.
2 AUDPC is area Ulder disease progress curve (% of disease days) from six late blight readilgs.
80
Research on Potato
en
~
__...30.--~~~~~~~~~~~~~
2500
-e
(1)
"'
2000
u"'
"#.
-o- Control
1500
500
<{
Treated
20
(;
.g
::::>
-a- Control
'6o-
&'15
~ 1000
o..
E' 25
O>
--Treated
cU
(1)
Kabete 1999 SR
Asante
Kabete 1999 SR
Asan te
10
1'--~...._~_._~__._~~..__~.....__~_,
30
20
40
30
50
60
40
Asan te
Asan te
en
>-
50
60
70
80
90
70
Loreto 1999 SR
Loreto 1999 SR
2500
cU
-e
(1)
"'cU
(1)
u"'
2000
1500
(;
~ 1000
.o 10
1-
o..
:::i
500
::::>
<{
30
o
20
40
30
cU
60
"'cU
(1)
u"'
2500
~ 1000
500
::::>
<{
90
2000
1500
o..
80
-e
(1)
50
60
70
Days after planting
70
Kalengyere 1999 SR
Victoria
en
>-
50
40
o
20
30
40
50
60
70
80
90
81
Asante
-tP
20
15
(i;
.o
10
~
o
:::J
Kabete 2000 SR
-o- Control
25
Dl
Treated
.,,.-
___ .-..........
..........
A.---------o----1.J---A,_.-
...... A
10'--~---'-~~....1-~~.L_.~--1.~~~
50
60
80
70
Asante
90
100
Loreto 2000 SR
30r-~~~~~~~~~~~~----.
~ 25
E
-~ 20
~
&' 15
(i;
.g
10
10'--_,c_--'--~--1...~~L_~...L~_L~_J
40
50
60
Victoria
70
80
90
100
Kalengyere 2000 SR
~30r-~~~~~~~~~~~~~
E 25
Dl
;
~
bo----...6.--~
20 ....
o 15 ,_
(i;
.o
10 ....
40
50
60
70
80
90
Days after planting
100
110
82
Research on Potato
Discussion
Applying the protective fungicide, Dithane
M-45, significantly reduced the development and severity of late blight and
increased tuber yields. In general, disease
was less severe in experimental plots with
the fungicide applied at intervals of seven
days, compared to the 21-day intervals or
control plots. The low disease levels in
Tigoni and Rutuku could be because both
varieties have relatively high levels of
horizontal resistance to the disease
compared to Asante/Victoria and bale.
In this study, a combined analysis of
variance was used for Asante/Victoria
because it was the only variety that was
planted in all cropping seasons. The lack
of consistency in disease levels recorded
across sites and seasons may be attributed
to environmental variation among sites. In
sorne cases, the early occurrence of the
disease befare the initiation of the spray
p~ogram cou Id account for madequate
d1sease control aRd confounding effects on
treatments. Adjustment of fungicide
Table 3. Combiled alalysis of varialce (ANOVA) Ol the effect of fulgicide applicatiol iltervals Ol late blight
severity ald tuber yield Ol AsalteMctoria variety (Victoria) plalted at Kabete, Loreto, ald Kalelgyere
sites durilg 1999 ald 2000 croppilg seasols.
So urce
DF
F-Value
Pr > F
Late blight severity
Ye ar
1
3.67
0.1956
2
Locatiol
1.25
0.04 a
2
Locatio l *year
118.36
0.0001 b
Rep (year*locatiol)
12
0.76
0.6887
Fulgicide 1
3
0.63
0.05 b
Fulgicide*year
0.97
3
0.4676
Fulgicide*locatiol
6
1.09
0.4613
Fulgicide*year*locatiol
42.68
0.0001 b
6
Fulgicide*rep (year*locatiol)
3.24
36
0.0001 b
Assess.day-DOA (year*locatiol)
28
93.77
0.0001 b
DOA*fulgicide (year*locatiol)
84
17.25
0.0001 b
Yield (t/ha)
Year
1
38.09
0.0001 a
Location
7.74
2
0.0006 b
Locatiol*year
29.76
2
0.0001 b
12
Rep (locatiol*year)
1.58
0.1424
Fulgicide
2.70
0.0466 a
3
3
Fungicide*year
0.47
0.7142
Fulgicide*location
0.60
6
0.7223
Fulgicide*location*year
8.25
0.0001 a
6
Rep*fulgicide (location*year)
12
0.30
0.9867
Tuber numbers
Ye ar
1
0.09
0.7933
2.20
2
Locatiol
0.3124
15.02
2
Locatio l *year
0.0005 b
12
Rep (location*year)
2.05
0.0481 a
Fungicide
1.79
3
0.2496
Fungicide*year
4.09
3
0.0673
Fungicide*location
2.18
6
0.1829
Fungicide*location*year
1.08
6
0.3942
Rep*fungicide (location*year)
0.96
36
0.5476
a = Significant at 0.05; b = Significant at 0.01.
1 Refers to application intervals of 7, 14, 21 days and control (no application) of the contact fungicide Dithane M-45.
83
{C)
{%)
{mm)
16
15
16
17
86
85
85
87
88.3
126.3
120.7
42.2
18
19
18
18
64
62
76
73
26.1
11.4
348.0
229.3
16
17
16
16
89
85
89
83
36.0
61.1
398.2
234.1
16
16
16
16
87
86
85
85
114.0
52.2
1.3
22.4
18
19
19
18
57
64
59
56
33.5
18.4
187.7
111.3
18
16
14
13
78
84
83
82
195.2
73.3
37.5
23.1
84
Research on Potato
Condusions
We conclude that applications of protective fungicide have a significant deterrent
effect on the development of late blight in
sorne sites in tropical Africa, and a
positive effect on accumulation of potato
biomass. The magnitude of disease
development and dry-matter partitioning
appears to be dependent on the levels of
resistance of the varieties and the frequency of fungicide applications.
However, further data are needed to assess
the contribution of cardinal factors such as
temperature, photosynthetic active
radiation, and rainfall in crop development
(tuber initiation, rate of tuber bulking) in
tropical Africa. Furthermore, elements
such as the effect of planting and harvesting dates on disease development and
potato biomass have not been adequately
addressed in the tropical highlands of
Africa.
References
Adipala, E. 1999. Potato production in
Uganda: A survey perspective. A report
to the Rockefeller Foundation by Dept.
of Crop Science, Makerere University,
Kampala, Uganda. 41 p.
Campbell, C.L. and L.V. Madden. 1990.
lntroduction to plant disease
epidemiology. John Wiley & Sons, NY,
USA.
El-Bedewy, R., O.M. Olanya, P.T. Ewell,
C. Lung'aho, P.S. Ojiambo, and
J. Karinga. 2001. Evaluation of potato
germplasm (populations A & B) for
85
86
Research on Potato
A potato (Solanum tuberosum L.) late blight (LB) simulation model was validated near Quito, Ecuador, at three sites and over two growing seasons. Two
varieties differing in resistance to Phytophthora infestans (Mont.) de Bary, the
causal agent of LB, were grown at each site. Data taken from each site
included hourly temperature and relative humidity and daily rainfall. Using
weather data from each site, the model simulated disease sever.ity for each
variety. Resistance parameters for the model were taken from measurements
made on the Peruvian varieties Amarilis-INIA (resistant) and Tomasa Tito
Condemayta (susceptible). At each site, the model simulated realistic differences between susceptible and resistant varieties. However, for the
experiments conducted where average daily temperatures were low (less
than 14C), the model underestimated disease severity. These temperatures
are characteristic of the principal potato growing area of Ecuador. The most
accurate simulation occurred at the Tola site in both seasons, where average
daily temperatures are above 14C. Overall, temperature was inversely
related to the predictive capability of the model.
Late blight is a major constraint to potato
production worldwide. The disease is
especially damaging in the tropical
highlands of Latin America, Asia, and
Africa, where it can be controlled only by
routine applications of fungicides, sometimes in conjunction with moderate levels
of host plant resistance (Forbes and Jarvis,
1994; Haverkort, 1990). For many poor
farmers, however, the disease is not
control led, losses are heavy, and crop
abandonment is not uncommon.
Resistant varieties are being developed for
the tropical highlands, but little is known
of how this resistance may best be used to
reduce fungicide application. General
recommendations for reductions in dosage
or for increases in the intervals between
fungicide sprays will probably be of only
partial benefit in the tropical highlands
1
87
88
Research on Patato
Table 1. Resistance components far three host resistance levels used in SAS version 2 of the late blight
simulator.
Resistance level 1
Latent period 2
lnfection efficiency3 Lesion expansion 4
Sporulation5
Tomasa (susceptible)
Yungay (moderately
resistant)
Amarilis (resistant)
63
1.0
4.10
257
63
0.9
3.66
228
76
0.8
3.40
107
1 Resistance
parameters were measured experimentally for three Peruvian varieties: Tomasa, Yungay, and Amarilis
(Andrade-Piedra Naranjo, 2000).
2 Base number in hours. Base number is modified in the model using the function latent period = base number /
temperature function, where temperature function = 0.0399* temperature + 0.0831 (Andrade-Piedra, 2000).
3 Relative values used to adjust weather-dependent functions.
4 lncrease in lesion diameter, mm/d.
5 Sporangia/mm 2/d.
Table 2. Research sites near Quito, Ecuador, where experiments were conducted in 1996/97 and 1997/98 to
validate a late blightsimulation model designed and validated previously in the temperate zone.
Site1
Altitude Average
Planting
Variety
Disease
First
Lesions/
(m)
T/RH 2
dates
evaluations
symptoms
plant
(no.) 4
(dd/mm/yy)
(dd/mm/yy) (dd/mm/yy) 3
CIP highland research
3060 11.4/11
18/12/96
Catalina 10/02/9710/02/07
10
station (CIP), 1996/97
Bolona
04/04/97
CIP highland research
3060 12.7/7
10
Catalina 30/12/975/11/97
22/12/97
station (CIP), 1997/98
Gabriela
30/03/98
Instituto Andino
2700 13.4/14 23/01/97
Catalina 06/01/9710
06/01/97
Superior de
Bolona
15/05/97
Agropecuaria (IASA),
1996/97
Catalina 13/04/9820
Pinantura (Pinantura),
3500 9.4/16
31/03/98
22/01/98
Uvilla
16/06/98
1997/98
2500 16.1/14 29/12/97
Catalina 17/02/98Centro Acadmico
04/02/98
Bolo na
Docente Experimental
21/04/98
La Tola (Tola),
1996/97
Catalina 02/02/9902/02/99
Centro Acadmico
2500 16.6/16 02/12/98
Uvila
13/04/99
Docente Experimental
La Tola (Tola),
1997/98
1 The
term in parentheses corresponds to reference to the site in the text and in Figure 1.
befare the slash = average daily temperature (T) in C during the trial; the number after = average hours/day
when relative humidity (RH) was ~ 90%.
3 Estimated date of first symptoms visible in the field. Oisease simulation was initiated one week befare this date.
4 Number of lesions/plant at the beginning of each simulation.
2 Number
89
Results
Field validation of the simulator
In ali six cases, the simulator predicted
slower disease progress for the resistant
variety (Figure 1). This demonstrates that
the model is fairly robust in its handling of
90
Research on Potato
Discussion
This study demonstrated that a patato LB
simulator designed for and validated in the
temperate zone has limited applicability
to tropical highland patato production. The
model simulated realistic differences in
host resistance between a resistant and
susceptible variety, but frequently underestimated disease severity for both.
Comparison of MU DPC with weather
factors indicated that underestimation was
associated with low temperatures characteristic of the main production zone in
Ecuador, between 3000 and 3500 m.
In this study we used SAS version 2 of the
model, which had been validated in Peru
in Comas and Oxapampa. Comas at 2400
m and Oxapampa at 1800 m represent the
lower fringe of patato production in the
central Andes. The validation studies
carried out in Peru demonstrated that the
model worked well for plots not treated
100
80
100
CIP,
1996 -1997
80
1 11
CIP,
1997 -1998
60
40
20
01 /15/1997
02/15/1997
03/15/1997
100
80
04/15/1997
IASA,
1996 -1997
12/1 0/1997
80
60
40
40
20
20
o
03/20/1997
04/20/1997
05/20/1997
100
80
02/20/1998
100
60
02/20/1997
01 /15/1998
Pinantura,
1997 -1998
li
"
Zl
.~
04/01/1998
04/15/1998
05/01/1998
06/01/1998
07/10/1998
04/01/1998
05/10/1998
100
TOLA,
1996 -1997
80
60
60
40
40
20
20
03/15/1997
04/15/1997
05/15/1997
06/15/1997
TOLA,
1997 -1998
02/01/1998
03/01/1998
Figure 1. Observed and simulated late blight severity far six experiments conducted near Quito, Ecuador. In each
plot salid squares represent observed disease severity of a susceptible variety; empty squares represent disease
severity of a resistant one. Salid lines represent simulated susceptible (highest line) and resistant varieties. Site
descriptions, varietal descriptions, and sorne parameters used far simulation are given in Table 2.
91
0.35 -
0.30 - o
0.20 (.)
a..
::>
<t:
0.25
0.15
rr
o
o
0.10 -
-<]
0.05 -
0.00 -
10
11
12
13
14
15
16
17
11
12
13
14
15
16
90%
Figure 2. Relation between MUDPC (a measure of model accuracy) and average daily temperature (A) and
hours per day of RH equal to ar above 90% (B). In each plot, salid squares represent susceptible varieties and
empty squares represent resistant varieties.
disease severity (data not shown). This
aspect of simulation does not appear to be
related to the problem of underestimation.
The resistance parameters we used in SAS
version 2 were derived from Peruvian
patato varieties (Table 1). Nonetheless,
when we simulated disease in Tola, where
the simulator appeared to work best, the
difference in resistance levels between
varieties Catalina and Gabriela was
accurately simulated using the parameters
of the Peruvian varieties Amarilis and
Tomasa, respectively. This occurred in two
trials. Even in IASA, where the simulator
underestimated disease severity, the
relative difference between the resistant
and susceptible varieties was well simulated using the Peruvian parameters. The
overall poor performance of the model in
the other experiments precludes this type
of evaluation, but it would be extremely
fortuitous if the parameters of Amarilis and
Tomasa are useful far many comparisons
between resistant and susceptible varieties. Eventually, the accuracy of the
resistance parameters used in the model
will probably depend on the intended use
of the simulator. Developing general
92
Research on Potato
References
Andrade-Piedra N., J.L. 2000.
Epidemiologa del tizn tardo de la
papa en los Andes peruanos: Validacin
del simulador LATEBLIGHT. MSc thesis.
Universidad Nacional Agraria La
Molina, Lima, Peru. 149 p.
Bruhn, J.A. and W.E. Fry. 1981. Analysis of
potato late blight epidemiology by
simulation modeling. Phytopathology
71 :612-616.
Doster, M.A. and W.E. Fry. 1991. Evaluation by computer simulation of
strategies to time metalaxyl applications for improved control of potato late
blight. Crop Protection 10:209-214.
Doster, M.A., M.G. Milgroom, and W.E.
Fry. 1990. Quantification of factors
influencing potato late blight suppression and selection for metalaxyl
resistance in Phytophthora infestans: A
simulation approach. Phytopathology
80:1190-98.
Fohner, G.R., W.E. Fry, and G.B. White.
1984. Computer simulation raises
questions about timing protectant
fungicide application frequency according to a potato late blight forecast.
Phytopathology 74:1145-47.
Forbes, G.A., X.C. Escobar, C.C. Ayala, J.
Revelo, M.E. Ordonez, B.A. Fry, K.
Doucett, and W.E. Fry. 1997. Population
genetic structure of Phytophthora
infestans in Ecuador. Phytopathology
87:375-380.
Forbes, G.A. and M.C. Jarvis. 1994. Host
resistance for management of potato
late blight. In: Zehnder, G., R. Jansson,
and K.V. Raman (eds.). Advances in
potato pest biology and management.
32:251-272.
Kato, M., E.S. Mizubuti, S.B. Goodwin,
and W.E. Fry. 1997. Sensitivity to
protectant fungicides and pathogenic
fitness of clonal lineages of
Phytophthora infestans in the United
States. Phytopathology 87:973-978.
Raposo, R. D.S. Wilks, and W.E. Fry. 1993.
Evaluation of patato late blight forecasts
modified to include weather forecasts:
A simulation analysis. Phytopathology
83:103-108.
Shaner G. 1973. Evaluation of slow
mildewing resistance of Knox wheat in
the field. Phytopathology 63:867-872.
Shtienberg, D., M.A. Doster, J.R. Pelletier,
and W.E. Fry. 1989. Use of simulation
models to develop a low-risk strategy to
suppress early and late blight in patato
foliage. Phytopathology 79:590-595.
Shtienberg, D. and W.E. Fry. 1990. Field
and computer simulation evaluation of
spray-scheduling methods for control of
early and late bl ight of pota to. Phytopathology 80:772-777.
93
andigena
E. Mihovilovich, L. Alarcn, A.L. Prez, and M. Bonierbale1
95
96
Research on Potato
primary infection 30 days after inoculation, and on mixed samples of leaves from
the two daughter plants during secondary
infection trials 50 days after tuber planting. Percentages of resistance were
estimated as the number of noninfected
plants (negative to ELISA), over the total
number of plants tested per family and
replication. Single plants were used to
represent genotypes within families. The
percentage of resistance on a family basis
was taken as a quantitative variable to
represent the mean progeny performance.
Data taken on percentage of resistance
were transformed to are sine for better
approximation of normal distribution.
General combining ability (GCA) and
specific combining ability (SCA) components of variance were estimated from the
expected mean squares of the analysis and
translated to genetic components under a
model of tetraploid inheritance assuming
random chromosome segregation (a = O),
no epistasis, no maternal effects, and
i ndependent assortment:
by
97
Results
Analysis of the diallel design
Table 1. Percentage of resistance to potato leafroll virus (PLRV) of 21 S. tuberosum subsp. andigena
families generated in a diallel mating design.
PPP-1161
LOP-868
Family
ZIM-440
CCC-4932
CUA-106
HJT-5535
HAW-5687
1O
33
20
43
40
83
13
ZIM-440
17
7
20
83
23
42
85
38
CCC-4932
27
35
PPP-1161
88
83
80
LOP-868
27
CUA-106
HJT-5535
Table 2. Analysis of variance for percentage of resistance of diallel mating design in S. tuberosum subsp.
andigena.
Df
MS
EMS
So urce
2
69.24" 5
Blocks
348.73**
20
Families
1091.56**
GCA
6
efe + cr2s + (p-1) cr 2g
SCA
14
40
Error
30.37lS
16.29
#e + efs
efe
Notes: MS = mean squares for are sine of percentage of non-infected plants in secondary infection trial; GCA = general
combining ability; SCA = specific combining ability; ** = Significant at the 1% level; ns = nonsignificant; CV = 17.3%.
98
Research on Potato
99
Table 5. Levels of resistance to PLRV infection determined far 7 parental accessions of the diallel design,
13 additional andigena clonal accessions, and 25 progenies from the parent accession LOP-868.
Accession
lnfection by clonal evaluation (%)
Reaction
Resistance in OP25 aphids/plant
50 aphids/plant
families1 (%)
Diallel parent accessions2
Highly resistant
88
o
LOP-868
O
nt 3
Susceptible
100
HAW-5687
80
nt
Susceptible
100
CCC-4932
100
nt
Susceptible
100
PPP-1161
100
nt
Susceptible
100
HJT-5535
100
nt
Susceptible
100
ZIM-440
100
Accessions
nt
Highly resistant
o
o
OCH-7643
Highly resistant
83
o
o
HUA-332
88
Resistant
50
o
VID-11
PPP DGV 94
20
90
Moderately resistant
76
CUP-199
90
100
Susceptible
47
OCH-11195
100
100
Susceptible
42
PPP SA 2103
100
100
Susceptible
29
PPP 1627
100
100
Susceptible
6
UNSAM 51
100
100
Susceptible
4
HMX 1686
100
100
Susceptible
4
CCC-4726
100
100
Susceptible
OCH-8225
100
100
Susceptible
nt
HJA 1489
100
100
Susceptible
nt
Clonal evaluation of 25 progenies from the family of the cross LOP-868 x HJT 5535
22 genotypes
O
O
Highly resistant
2 genotypes
O
10
Highly resistant
1 genotype
20
70
Moderately resistant
Controls
DW.84-1457
O
20
Highly resistant
nt
Achirana INTA
30
80
Moderately resistant
nt
Perricholi
100
100
Susceptible
nt
1 OP = open pollinated.
2 Parent accession CUA-106 was not tested because it was not available in virus-free in-vitro stocks.
3 nt = not tested (open pollinated seed not available in germplasm stocks).
100
Research on Potato
Discussion
Resistance to PLRV in the commercial
patato tuberosum has proved difficult to
manipulate in breeding due to the nonabsolute and complex genetic nature of the
resistance. For this reason, there are few
commercial varieties with substantial
resistance to PLRV infection (SolomonBlackburn and Barker, 1993). Levels of
resistance high enough to avoid infection
have been reported in wild Solanum
species (Hooker, 1977). Special measures,
however, are often necessary to overcome
crossing barriers, in addition to the significant ti me and effort requ i red to i ntrogress
resistance from unadapted germplasm into
agronomical ly acceptable varieties.
Andigena is a native cultivated species
genetically similar to the commercial
patato due to their clase evolutionary
origins (Estrada, 1999). This explains the
ease of crossability between the two
subspecies, whereas the divergent selection that has been applied to them in
recent history accounts for the heterosis
found in sorne of their hybrids (Tarn and
Tai, 1983). These attributes and those
described befare, make andigena a
desirable genetic source for patato
breeding.
92.111 x C93.139
40
(Susceptible control)2
1 Mean of three replications.
2 Cross between two PLRV-susceptible breeding clones
generated at CIP.
101
102
Research on Potato
Acknowledgments
We are gratefu 1 to R. Rivera, F. Sagu ma,
and A. Gmez for their support in greenhouse and field experiments, and to
Dr. H. Mendoza for consultation on
statistical analysis.
References
Bastos, C. 1997. Resistencia en clones de
germoplasma de papa (Solanum sp.) al
PLRV, PVS y APMV. Ingeniero
Agrnomo thesis. Universidad Nacional
del Centro del Per. Huancayo, Peru.
43 p.
Brown, C.R. 1980. lncorporation of virus
resistance and future plans. In: Report
of the planning conference on the
strategy for virus management in
potatoes 11 held 21-25 Apri 1 1980 at the
103
105
106
Research on Potato
107
.......
oCX>
~
,.,
(J>
(!>
g.
o
::::1
a"
Table 1. Wilt incidence, yield, and percentage infected tubers (visible and latent) of advanced patato clones plantad in a race 3/Biovar 2A-infested field in Carhuaz, Peru,
selected far their high to moderate resistance to bacteria! wilt in 1998-1999 and 1999-2000 crops.
1999 Planting
1998 Planting
Wilted
Total
Rotted tubers Infectad
Wilt
Wilted
Total
Pedigree
Wilt
Rotted tubers Infectad
plants
at harvest
incidence
plants
Yield
tubers
incidence
Yield
tubers
at harvest
(%)
(% weight)
(%)
(g/plant)
(%)
Score
(%)
(g/plant) (% weight)
seo re
o
o
o
1 (BWH87.344R X TXY.11 )1
1048
1.00
858
27.0
1.00
3
o
1268
903
31
1.00
o
36.4
2 (CRUZA-148 X C90.205)8
1.00
5
o
o
o
1.00
1264
18.5
1.00
702
31.6
8
3 (720118.1 X C90.205)21
o
1378
19.2
1.00
o
672
24.7
1.00
6
o
4 (720118.1 X C90.205)20
o
o
19.2
1.00
o
1556
1.00
860
5
30.7
5 (720118.1 x BWH87.183)3
o
1518
o
36.1
1.04
1320
1.00
1
1
40.3
6 (720118.1 X C90.205)9
o
o
1.04
1.00
460
1
745
1
38.4
7 (C90.205 X 34.73)13
o
o
1.04
1270
1.00
680
1
37.1
8* H89.65.44
3
o
o
1375
32.5
1.12
728
1.00
3
2
31.8
9 (BWH87.415 x DXY-7)16
o
690
9
1.16
4
356
23.1
10 (34.73xTXY.11)2
1.00
4
o
o
893
14.1
1.16
4
1430
33.6
11 (CRUZA-148 x BWH87.344R)1
1.00
5
o
1834
13
9.1
1.16
4
1420
12 (BWH87.230R x C90.205)1
1.00
1
31.1
o
750
4
1.20
948
39.8
13 (720118.1 X C90.205)17
1.00
5
3
o
15
270
22.5
1.20
522
60.7
14 (34.73 x BWH87.344R)1
1.00
5
5
o
7
29.5
1.24
1466
30.7
1.00
1136
6
6
15 (CRUZA-148 x BWH87 .344R)3
o
o
60.0
1.28
7
700
1.00
510
1
18.8
16 (28.68 X C90.205)5
o
o
1.28
20.6
1.00
880
7
598
5
17* 392289.34
1.28
o
950
5
10.4
7
1260
4
30.0
18 (28.68 x BWH87.344R)13
1.00
o
o
1.28
7
1020
675
3
19.0
19* BWH87 .338
1.00
o
o
1840
1.32
1104
53.1
1.00
8
3
20 69.4
o
1972
1
1.32
1060
57.0
1.00
8
4
21 (CRUZA-148 X C90.205)12
o
o
57.1
1.00
136
22 (BWH87.215 X XY.13)106
o
1056
2
17.1
1.00
23 (BWH87.409xXY.16)15
o
o
45.8
1.00
2325
24 23.3
o
1
68.2
1.00
1980
25 (5.63 x BWH87.183)5
2
o
28.6
26 (5.63 x BWH87.344R)8
1.00
560
o
o
16.7
1.00
700
27 {5.63 X C90.205)2
o
6
19.0
1.00
530
28 (28.68 X C90.205)7 '
o
7
52.0
1.00
725
29 (720118.1 X C90.205)2
C")
=ti
"'O
co
;:
3
~
-e
o
;::::.
e.o
e.o
e.o
1
NI
o
o
o
tO
30 (TXY-8 x BWH87.183)16
1.00
31 (TXY-8 x BWH87.344R)6
1.00
32 (BWH87.172R xTXY-2B)12
1.00
33 (BWH87.344R X TXY.11)18
1.00
34 (381064.2xTXY-11)4
1.00
35 (TXY.11 x BWH87.344R)1
1.00
36 (BWH87.420 X C90.205)6
1.00
37* SR 17.50
1.00
1.00
38* 392278.19
39 (BWH87.446R X TXY.2B)1
1.00
40 (720118.1 x BWH87.183)6
1.00
41 (BWH87.407R X TXY.11)1
1.00
42 A1-12.55
1.00
43 (C91.640 X 34.73)2
1.00
44 (BWH87.289xXY-13)103
1.00
45 (381064.2xTXY-11)1
1.00
46* BWH87 .176 x XY.16)24
1.00
47* 392270.2
1.00
48 (720118.1 x BWH87.183)5
1.00
49 (BWH87.230R X C90.205)5
1.00
50 (720118.1 x BWH87.183)1
1.00
51 * 392285.72
1.00
1.00
52 (3.5 X C90.205)3
53 (BWH87.230R X TXY.11)3
1.00
54 (BWH87.528R X TXY.11)1
1.00
55 (5.63 x BWH87.183)1
1.00
56 (TXY-8 x BWH87.183)8
1.00
57 (28.68 x BWH87.344R)6
1.00
Yungay (S) t
2.83
Revolucion (S)
4.36
Molinera (MR) t
3.18
Cruza 148 (R} t
1.15
* Clones selected from the 1997/98 evaluation trial.
o
o
o
o
o
o
o
o
o
o
o
o
o
o
o
o
o
o
o
o
o
o
o
o
o
o
o
o
45.7
84.2
54.5
4.0
1928
323
1340
372
1444
620
1494
1320
740
914
3
4
9
41
2
616
633
780
517
250
367
480
1340
1700
1580
767
950
925
460
1540
1300
880
628
1094
870
1027
25.2
35.4
43.4
60.0
o
4
o
o
5
o
o
52
o
o
o
o
22
o
o
21
12
30
6
2
10
21
50
15
3
47.1
9.5
34.2
1.40
1.40
1.40
1.48
1.48
1.48
1.56
1.64
1.68
1.68
1.68
1.68
1.80
1.84
1.88
1.88
1.92
2.04
10
10
10
12
12
12
14
16
17
17
17
17
20
21
22
22
23
25
1428
826
809
564
456
521
3
1
7
3
4
13
584
507
920
920
656
686
558
1362
676
1264
1018
4
4
2
4
2
6
5
2
4
7
11
2.08
2.12
2.20
2.20
26
28
30
30
660
1178
2080
1314
2
21
23
10
33.8
41.5
42.6
3.78
4.66
2.36
1.24
70
92
33
6
1106
874
862
1349
34
46
27
12
19.5
72.8
50.0
24.8
10.3
20.0
23.5
36.6
14.6
37.0
66.3
40.4
38.5
33.3
45.5
9.0
21.2
71.3
71.5
27.6
42.2
Laboratory trials
Tubers were washed in tap water and
disinfected by dipping them in 1% sodium
hypochlorite for 1O min. They were then
graded in three size categories: 1.5-2.5
cm, 2.6-5 cm, and > 5 cm. Tubers were
then tested individually for latent infection
by R. solanacearum using the CIP
postenrichment NCM-ELISA kit as described by Priou et al. (1999a, 1999b). A
total of 2246 tubers were tested in 1999
and 4329 tubers in 2000. Upan cutting the
stolon end of the tuber to remove the
vascular ring for testing in ELISA, tubers
that exhibited visible oozing were not
further processed, and the percentage of
tubers showing visible symptom was
scored per clone. In 1999, the enriched
samples were streaked on modified
Kelman's medium (French et al., 1995) to
confirm the presence of the bacterium.
When ELISA was positive but the bacteria
could not be isolated, DNA amplification
by PCR was performed from enriched tuber
extracts (Priou et al., 2001 ).
Statistical analyses
Qualitative assessment
Tables of probabilities were calculated
according to the binomial distribution to
determine the probabilities of finding at
least one infected tuber in a total number
of sampled tubers per clone. This probability is:
1 - (1 - p)K
11 O
Research on Potato
where:
P is the probability of finding one
i nfected tu ber.
(1-p)k is the probability of finding no
infected tubers in k tubers.
K is the number of sampled tubers per
clone for analysis, e.g., the number of
tubers sampled for each plant multiplied
by the number of plants per clone.
The main assumptions are that the disease
spreads uniformly throughout the field,
each plant (of a given clone and plant
infection leve!) has the same probability of
being infected at the same rate of infection, and the probability of any tuber to be
infected is the same, independent of the
plant.
Quantitative assessment
Statistics for estimation of sample size for
proportions based on the normal distribution have been used to determine the
necessary number of tubers/clone to
sample at random to accurately estmate
the true frequency of infected tubers that
would be obtained by processing ali tubers
of the clone. The estimation avoids having
to process al 1 tubers/clone. The sample
size n depends on the precision leve!
required d, the infection rate per clone p,
and the population size N = total number
of tubers for the clone in the field:
n = n0 /f
111
Quantitative assessment
Statistical analysis was appl ied to determine the optimum number of tubers to
sample per clone that al lows an accurate
estimation of the proportion of infected
tubers of the clone. Sample sizes (n) to be
analyzed in ELISA were calculated with
proportions of infected tubers (p) (at
various levels: minimum, average, and
maximum obtained from the 35 clones in
the 1999 trial) and precision levels (d),
according to the total number (N) of tubers
harvested per clone (depending on the
number of plants and on the multiplication
rate of the clone). The average number of
tubers/clone from the harvest of 1O plants
in the 1999 trial was 124 (12.4 tubers/
plant). Precision levels have been chosen
according to their significance for various
infection rates, and to limit the sample
Table 2. Average infection rates/plant in relation to the average tubers/plant far CIP advanced clones
separated in two groups according to the proportion of infected tubers (results obtained in 1998/99
and 1999/2000 evaluation trials).
Group 1
Group 2
~ 30% infected tubers
> 30% infected tubers
lnfection rate/plant (av)
lnfection rate/plant (av)
1998 planting
1999 planting
1998 planting
1999 planting
No. tubers/plant (av)
0.30
0.20
0.40
0.40
10
0.20
0.20
0.45
0.50
20
0.50
0.53
0.20
0.23
30
112
Research on Patato
Table 3. Probabilities to obtain at least one infected tuber from (x) tubers sampled (e.g.,
5, 1O, 15, 20, or 25 plants according to the clone infection rate (IR).
Probability using 5 plants
IR
(x=1)
(x=2)
(x=3)
(x=4)
(x=5)
(x=6)
(x=7)
0.1
0.410
0.651
0.794
0.878
0.928
0.958
0.975
0.2
0.893
0.672
0.965
0.988
0.996
0.999
1
0.3
0.832
0.972
0.995
0.999
1
1
1
0.4
0.922
0.994
1
1
1
1
1
0.5
0.969
0.999
1
1
1
1
1
1
0.6
0.990
1
1
1
1
1
Probability using 10 plants
IR
(x=2)
(x=3)
(X=1)
(x=4)
(x=5)
(x=6)
(x=7)
0.1
0.651
0.878
0.958
0.995
0.985
0.998
0.999
0.2
0.893
0.988
0.999
1
1
1
1
1
1
1
1
0.3
0.972
0.999
1
1
0.4
0.994
1
1
1
1
1
1
0.5
1
1
1
1
0.999
1
1
0.6
1
1
1
1
1
1
Probability using 15 plants
IR
(x=1)
(x=2)
(x=3)
(x=4)
(x=5)
(x=6)
(x=7)
0.991
0.1
0.794
0.958
0.998
1
1
1
1
1
0.2
0.965
0.999
1
1
1
1
0.3
0.995
1
1
1
1
1
1
0.4
1
1
1
1
1
1
Probability using 20 plants
IR
(X=2)
(x=4)
(x=5)
(x=6)
(x=7)
(x=1)
(x=3)
0.998
1
0.1
0.878
0.985
1
1
1
1
1
1
1
1
0.2
0.988
1
1
1
1
0.3
1
1
1
0.999
1
1
1
1
1
1
0.4
1
Probability using 25 plants
(x=6)
(x=7)
(x=4)
(x=5)
(x=3)
(x=1)
(x=2)
IR
1
1
1
1
1
0.928
0.995
0.1
1
1
1
1
1
1
0.2
0.996
1
1
1
1
1
1
1
0.3
1
1
1
1
1
1
0.4
1
sizes to reasonable numbers. For instance,
for clones having an infected tuber rate of
0.4-0.6, a precision of 0.15-0.20 is
sufficient. This proportion would quickly
increase after harvest in any case. But for
genotypes having only a 0.1 proportion of
infected .tubers, the precision should be
higher (< 0.1) to avoid having negative
detection and rating the clone as resistant.
lf 1 O plants/clone are to be assessed,
analyzing 30 tubers/clone provides an
estimation of the proportion of infected
tubers with a precision that varies between
0.095 and 0.15 for various infection levels
(x=B)
0.985
1
1
1
1
1
(x=9)
0.991
1
1
1
1
1
(x=B)
1
1
1
1
1
1
(x=9)
1
1
1
1
1
1
(x=B)
1
1
1
1
(x=9)
1
1
1
1
(x=B)
1
1
1
1
(x=9)
1
1
1
1
(x=B)
1
1
1
1
(x=9)
1
1
1
1
113
Table 4. Sample sizes (n) to analyze in ELISA to estimate the proportion of infected tubers according to the
total number of tubers N harvested per clone, far various infection levels.
Proportion p of
Precision level
Number (n) of tubers to be sampled according to the
infected tubers
d = 1Est. p - p 1
total number (N) of tubers per clone1
of the clone
(N=100)
(N=200)
(N=400)
(N=600) (N=lnfinite)
Minimum = 0.1
0.05
59
83
104
113
139
0.095
29
33
36
37
39
0.10
45
58
68
72
81
Average = 0.3
0.15
27
31
34
34
36
0.15
30
35
38
39
41
Maximum = 0.6
0.20
20
22
23
24
24
1
n = nclf where n0 = (t2pq)/d 2 and f = 1+(nofN). The Student's t value ata 95% confidence leve! was used.
Table 5. Precision levels (d) obtained after simulation of 15 random samplings of (n) tubers to be analyzed
in ELISA to estimate the proportion (p) of infected tubers far the 35 clones selected in 1999-2000 .
Number n of tubers to be
(n=25)
(n=30)
(n=35)
(n=40)
Probability
analyzed per clone
True p
Estimated p
0.3390
0.3389
0.3386
0.3385
0.3414
Precision level: d = 1Est. p - p 1
0.0024
0.0025
0.0028
0.0029
lower than the theoretical level of 0.1 5
shown in Table 4. Thus, analyzing 30
tubers per potato genotype when 1 O plants
are assessed gives an accurate estimate of
the proportion of infected tubers.
Furthermore, according to the results
obtained with the qualitative assessment
of sample size, this sample size would
ensure detection of tuber infection even if
the clone real infection rate were low
(equal or less than 0.1) ata probability
ranging from 95% to 100% (Table 3), so
that the clone could not be misevaluated
as resistant. For each clone, 30 tubers can
be taken at random from the heap of al 1
harvested tubers or three tubers per plant
can be randomly sampled from 1O plants
or two tubers from 15 plants.
Thus in a trial evaluating 20 clones with
approximately 300 tubers each harvested
from 25 plants, the number of tubers to be
processed would be 600 instead of 6000, if
ali tubers were to be analyzed. That would
save a lot of labor and supplies costs, and
moreover more tubers would remain for
other analysis (quality, storage).
114
Research on Potato
Conclusion
Breeding for resistance to BW at CIP has
resulted in a high to moderate level of
resistance to wi lt, superior to the phureja
x tuberosum hybrids. However, a high
frequency of latent infection is still present
in tubers at levels comparable to those
observed for S. phureja-derived clones,
such as Molinera, or for Cruza 148, one of
the less susceptible varieties. Ciampi et al.
(1980) found that sorne phureja x
tuberosum hybrids did not harbor tuber
latent infection by R. solanacearum race 3
when grown under cool conditions in the
greenhouse, even after incubation of
clones for 70 d at 28C. Anguiz and
Mendoza (1997) also reported the absence
of latent infection in sorne phureja-derived
clones after growing them in a race 1infested field. The detection procedure
they used (isolation on TZC medium),
however, lacked sensitivity compared with
postenrichment NCM-ELISA, which is 100to 1000-fold more sensitive (Priou et al.,
1999a). Probably, resistance to latent
infection does not occur in the genetic
base of the parents used for the crosses
that led to these materials. Latent infection in tubers was not a selection criterion
References
Anguiz, R.J. and H.A. Mendoza. 1997.
General and specific combining
abilities for resistance to bacteria! wilt
(Pseudomonas solanacearum E.F. Smith)
57:319-329.
p. 381-385.
Grimsley, N. and J.-F. Wang. 1994. Chair's
perspective: Host resistance. In:
Hayward, A.C. and G.L. Hartman (eds.).
Bacteria! wilt: The disease and its
causative agent, Pseudomonas
solanacearum. CAB lnternational,
Wallingford, UK. p. 197-199.
Kloos, J.P. and B.B. Fernandez. 1986.
Evaluation of patato germplasm for
resistance to Pseudomonas
solanacearum (E.F. Smith) and
adaptation in Mindanao. The Philippine
Agriculturist 69:263-276.
115
116
Research on Potato
The effect of Bt-cry1/a1 (cryV now designated cry1/a1 under revised nomenclature) of potato transgenic plants on the two species of potato tuber moth,
Phthorimaea operculella (Zeller) and Symmetrischema tangolias (Gyen), was
tested in Peru. Detached leaf bioassays were done using 1 O neonate larvae
per replication on each transgenic line of bred potato varieties Atlantic and
Spunta. Mortality in Atlantic transgenic plants was lower for P. opercule/la,
ranging from 18 to 34%, than for S. tangolias ranging from 40 to 94%. All
transformed Spunta lines tested showed high levels of mortality in both species, with mortality ranging between 80 and 98%. The Bt-cry1/a1 gene offers
another source of resistance that can be pyramided for effectiveness toward
the development of durable resistance to PTM and other insect pests.
117
Leaf bioassay
A detached leaf bioassay, developed by
Westedt et al. (1998), was used to test the
mortality of PTM feeding on Bt-cry7 Ja 7
transgenic plants. The petiole of the leaf
was i nserted i nto a sponge fitted in the top
of a glass vial full of water. The leaf and
the vial were then placed on a filter paper
disk in a Petri dish (25 x 150 mm). Ten
neonate larvae were introduced into each
Petri dish. Each Petri dish was considered
as a replication and five repl ications per
transgenic line were used in a completely
randomized design.
Table 1. Transgenic patato lines evaluated far control of patato tuber moth bioassays and their Bt-cry1/a1
gene constructs.
Vector
Construct
Line
pBlcry5
35S/Bt-cry1Ja1 /GUS
All Atlantic-Bt lines
pBIML5
35S/Bt-cry1Ja1
Spunta G-2
pBIML5
35S/Bt-cry1la1
Spunta G-3
pBIML 1
GSP/Bt-cry1/a1
Spunta S-1
GSP/Bt-cry1/a 1
pBIML 1
Spunta S-4
35S/Bt-cry1Ja1 /35S/PW
pBIML6a
Spunta 6a-3
118
Research on Patato
Statistical analysis
All mortality data were subjected to
arcsine transformations befare analysis.
Data from the two varieties were analyzed
separately. Means were compared using
the Waller-Duncan test at 0.05. SAS/STAT
software for Windows (SAS lnstitute 1989)
was used for statistical analysis.
Tuber bioassay
Antibiosis effect was tested in the laboratory by putting tubers and PTM larvae in
closed containers (no-choice test). Harvested tubers from Atlantic and Spunta
lines were divided into three groups of 50
g each (representing three replications) for
P. operculella tuber assays. Each replication (50 g of tuberlets) was exposed to 1O
neonate larvae.
For the Symmetrischema tangolias tuber
test, 30 g for Atlantic and 25 g for Spunta
were used. Each replication was exposed
to six neonate larvae for Atlantic and five
neonate larvae for Spunta. One PTM
neonate larva was used for each 5 g of
tuber weight. After 25 days, the number of
pupae developed was recorded and overall
mortality was expressed in percentage.
Tests were carried out in an insect massrearing room under controlled temperature
Table 2. Mortality and larval length reduction in patato tuber math, Phthorimaea operculella, in leaf and
tuber biaassay of Bt-cry1/a1 Atlantic and Spunta patato lines, La Malina, Peru, 2000.
Transgenic lines
Leaf assay1
Tuber assay1
Mortality (%)
Reduction (%)
Mortality (%)
Atlantic Bt-6
18 ab
65.9 a
27 ab.
Atlantic Bt-5
22 ab
65.7 a
40 a
Atlantic Bt-2
30 a
61.0 a
20 b
Atlantic Bt-4
34 a
66.3 a
30 ab
Atlantic (control)
5b
Oc
Spunta G-2
80 b
46.9 b
100 a
Spunta 6a-3
86 ab
63.3 b
100 a
Spunta G-3
90 ab
49.8 b
100 a
Spunta S-1
92 ab
46.6 b
100 a
89.5 a
Spunta S-4
98 a
100 a
Spunta (control)
ob
Oe
1 Means within a column followed by the same letter are not significantly different (P
= 0.05) by the
Waller-Duncan test.
GIP Program Report 1999 - 2000
119
Table 3. Mortality and larval length reduction in patato tuber moth Symmetrischema tango/ias in leaf and
tuber bioassay of Bt-cry1/a1 Atlantic and Spunta patato lines, La Malina, Peru, 2000.
Transgenic lines
Leaf assay1
Tuber assay1
Mortality (%)
Reduction (%)
Mortality (%)
Atlantic Bt-6
40 b
76.0 a
72 a
Atlantic Bt-4
56 b
71.4 a
78 a
Atlantic Bt-5
92 a
52.3 b
53 a
Atlantic Bt-2
94 a
85.7 a
50 a
Atlantic (control)
46.67 b
44 a
Spunta G-2
80 b
77.8 a
100 a
Spunta 6a-3
82 ab
77.4 a
100 a
Spunta S-1
90 ab
100 a
Spunta G-3
96 a
83.7 a
100 a
Spunta S-4
96 ab
87.9 a
100 a
Spunta (control)
1Oe
16 b
1 Means within a column followed by the same letter are not significantly different (P = 0.05) by the
Waller-Duncan test.
- = Data not available because surviving larvae escaped befare they could be measured.
Larval development was severely affected
on both transformed cu ltivars. The reduction in larval length ranged from 52.3 to
85.7% on AtlanHc and from 77.4 to 87.9%
on Spunta lines.
Of the PTM grown on Atlantic transgenic
lines, Symmetrischema tangolias larvae
were general ly more affected than were
P. operculella larvae.
Tuber bioassay
Mortal ity of P. operculella on Atlantic
transgenic lines was significantly different
than on the nontransformed control (Table
2). None of these lines gave 100% mortality in leaf and tuber bioassays. lnsect
mortalities ranged from 20 to 40%. This
level of larval mortality was lower than
the levels reported by Mohammed et al.
(2000) using sorne transgenic Atlantic lines
(Bt-2, Bt-4, and Bt-6) with stored and
newly harvested tubers.
All Spunta transgenic lines showed high
levels of r~sistance (100% larval mortality). These 'results are similar to the leaf
120
Research on Potato
Condusions
Mortal ity of both P. operculella and
Symmetrischema tangolias was found to
be high on leaves of transgenic Spunta
lines. In general, the larvae that fed on
Atlantic and Spunta transgenic plants
containing the Bt-cry1la1 gene were
-severely restricted in growth (reduction in
larval length in the leaf bioassay).
The res u lts of both PTM bioassays demonstrated that h igh leve Is of Bt-cry1Ja1
expression can be achieved with the genes
construct and vectors used in Spunta
transgenic 1 ines.
References
Barton, K. and M. Miller. 1993. Production
of Bacillus thuringiensis insecticida!
proteins, In: Kung, S. and R. Wu (eds.).
Transgenic plants. Vol.1, Engineering
and utilization. Academic Press, San
Diego, CA, USA. p. 297-315.
Bauer, L.S. 1995. Resistance: A threat to
the Insecticida! crystal proteins of
Bacillus thuringiensis. Florida Entomologist 78:414-443.
Caedo, V., J. Benavides, A. Golmirzaie,
F. Cisneros, M. Ghislain, and
A. Lagnaoui. 1999. Assessing
Bt-transformed potatoes for patato tuber
moth, Phthorimaea operculella (Zeller),
management. In: lmpact on a changing
world. Program Report 1997-98. lnternational Patato Center, Lima, Peru.
p. 161-170.
Crickmore, N., D.R. Zeigler, J. Feitelson,
E. Schnepf, J. Van Rie, D. Lereclus,
J. Baum, and D.H. Dean. 1998. Revision of the nomenclature for Bacillus
thuringiensis cry genes. Microbial and
Molecular Biology Review 62(3):807813.
Douches D., A. Westedt, K. Zarka,
B. Schroeter, and E. Grafius. 1998.
Patato transformation to combine
natural and engineered resistance for
controlling tuber moth. HortScience
33(6):1053-1056.
121
123
124
Research on Patato
7.-----------.-,...- .
,_..'
Y=0.677x+07614
~.-
_,/;~::./
:o
e 5
a.
....
~~
0.6
. O.O
-0.6
:~ : t
-2.4
gi
-2.9
...J
~.5
-4.1
...,.,,,-'
...
3 '---.,.-----.------.--...---_J -4.7
5
6
7
8
Log concentration (granules/mi)
Table 1. Linear regression analysis of irradiation time and log activity ratio of PoGV and its calculated halfinactivation time (t1h = log 0.5/s/ope).
Exp. ~adiat~on
Slope
t1h (95%
R2
lntercept
F (df)
Probability
No. mtens1ty
( St. err.)
confidence
( St. err.)
(W/m 2)
limits) (min)
1
820
-0.066 (0.021) . 4.54 (2.82 - 11.6) 0.675
0.62 (0.59) 10.38 (1, 5)
0.0234
2
790
-0.067 (0.010) 4.51 (3.52 - 6.26) 0.754
0.98 (0.29) 48.92 (1, 16) 0.0000
3
111 o
-0.115 (0.032) 2.62 (1.70 - 5.69) 0.653
0.29 (0.45) 13.16 (1, 7)
0.0084
-0.51 (0.29) 12.10 (1, 8)
4
700
-0.046 (0.013) 6.56 (4.19 - 15.03) 0.602
0.0083
y - log activity ratio; x - exposure time in minutes.
125
1b
1a
Jan. 2000: -820 W/m 2
~
:;;
~
Cl
3
-1
-2
-3
-4
75
50
time(min)
25
100
125
2a
2000
4000
energy (x 1000 joule)
6000
2b
Mar. 2000: -790 W/m 2
">
g>
...J
..,, -
-1
~
~
-2
-3
-4
50
25
75
time(min)
-3
-4'-----.......------------__.
100
125
2000
4000
energy (x 1000 joule)
6000
3b
3a
2
2 . - - - - - - - - Dec. 2000: -111 OW/m
">
n
as
g>
...J
-1
-2
-3
-.d.
25
75
50
time(min)
100
125
2000
W/m 2
6000
2.------------------,
--.-~
4b
4a
2 . - - - - - - - - - J a n . 2001: -700
4000
1st segment
2nd segment
as
e
ofl-1~
Cl
-2
-3
-3
4'----..,---..,----r------r---~
25
50
75
time (min)
100
125
-4L-----------..----....---'
2000
4000
energy (x 1000 joule)
6000
Figure 3. Bisegmented inactivation curves of PoGV during four series of exposure conducted under natural solar
radiation in Lima, Peru. Log activity is plotted against exposure time (1-4a) and the accumulated energy sum
calculated from measurements with a UCor 200S pyranometer,400-1100 nm (1-4b).
126
Research on Potato
Table 2. Linear regression analysis of radiation energy (400-1100 nm) and lag activity ratio of fbGV and
its calculated half-inactivation to energy (energyVz = lag 0.5/slope).
Radiation
Slope
t11z (95%
Exp.
lntercept
R2
intensity
( St. err.) confidence limits)
F (df)
Probability
No.
(
St. err.)
2
3
3
(W/m )
(x 10 )
(x 10 joule)
1
820
-1.34 (0.42)
225 (139 - 581)
0.672
0.61 (0.59) 10.23 (1, 5)
0.0240
2
790
-1.3 (0.18)
231 (180 - 320)
0.757
0.93 (0.28) 49.70 (1, 16)
0.0000
3
1110
-1.73 (0.48)
174 (113 - 379)
0.653
0.29 (0.45) 13.18 (1, 7)
0.0084
4
700
-1.06 (0.31)
284 (181 - 656)
0.599 -0.52 (0.29) 11.96 (1, 8)
0.0086
y = lag activity ratio; x = accumulated energy in joule.
Table 3. Polynomial
Exp. Radiation
No. intensity
(W/m 2)
1
820
2
790
3
1110
4
700
lt was not possible to explain the inactivation solely by the radiation energy sum.
However, a LiCor 2005 pyranometer only
approximates sunlight energy because of
its relative response along the energy
distribution in the solar spectrum. In
natural sunlight, 3.5-4% of the energy is
in the UV spectrum (300-400 nm). In all
experiments, the energy measured with
the UV-B sensor was only 0.2% of the
energy compared with the pyranometer.
Further experiments at different altitudes
should enable a comparison of total
energy and 1ight spectra.
Conclusions
The method employed is adequate to
calculate the half-inactivation time of
baculoviruses under natural radiation. lt
seems important to differentiate between
irradiation time and irradiation energy sum
during exposure. In recent investigations,
we started to test the persistence of PoGV
activity at various altitudes of the Peruvian
Andes. Additionally, the method is used to
evaluate the protective capacity of
formu lations.
regression analysis of radiation energy (400-1100 nm) and lag activity ratio of fbGV.
Constants of the polynomial model
F (df)
Probability
y = m1 x2 + m2x + b ( St. err.)
1.24
2.1
4.53
4.51
(0.45)
(0.51)
(1.70)
(2.11)
-1.41
-1.76
-2.32
-1.78
(0.30)
(0.27)
(0.58)
(0.55)
0.54
1.07
0.39
-0.35
(0.39)
(0.29)
(0.41)
(0.29)
0.862
0.806
0.732
0.681
37.38
53.86
17.73
11.72
(1,
(1,
(1,
(1,
12)
26)
13)
11)
0.0001
0.0000
0.0010
0.0057
127
Acknowledgment
This work is a collaborative project
between the University of Hohenheim,
Germany; Federal Biological Research
Center, Germany; and CIP, financed by the
Federal Ministry of Co-operation and
Development, Germany.
References
David, W.A.L. 1969. The effect of
ultraviolet radiation of known
wavelength on a granulosis virus of
Pieris brassicae. Journal of Economic
Entomology 14:336-342.
Finney, J.R. 1971. Probit analysis (Third
edition). Cambridge University Press,
UK. 333 p.
Griego, V.M., M.E. Martignoni, and
A.E. Claycomb. 1985. lnactivation of
nuclear polyhedrosis virus (baculovirus
subgroup A) by monochromatic UV
radiation. Applied and Environmental
Microbiology 49(3):709-71 O.
Huber, J. and C. Ldcke. 1996. UVinactivation of baculovirus: The
bisegmented survival curve. IOBC/
WPRS Bulletin 19(9):253-256.
lgnoffo, C.M., C. Garca, and
S.G. Saathoff. 1997. Sunlight stability
and rain-fastness of formulations of
Baculovirus heliothis. Environmental
Entomology 26:1470-1474.
Jones, K.A., A. Westby, P.J.A. Re 1ly, and
M.J. Jeger. 1993. The exploitation of
micro-organisms in the developing
countries of the tropics: In Jones,
128
Research on Patato
68:385-391.
McGuire, M.R., R.W. Behle, H.N. Holly,
and T.C. Fry. 2000. Calibration of a
sunlight simulator for determining solar
stability of Bacillus thuringiensis and
Anagrapha fa/cifera nuclear
polyhedrovirus. Environmental
Entomology 29:1070-1074.
Reed, E.M. and B.P. Springett. 1971.
Large-scale field testing of a granulosis
vi rus for the control of the potato moth
(Phthorimaea operculella (Zell.) (Lep.
Gelechiidae)). Bulletin of
Entomological Research 61 :223-233.
Shapiro, M. 1985. Effectiveness of B
vitamins as UV screens for the gypsy
moth (Lepidoptera: Lymantri idae)
nucleopolyhedrosis virus. Environmental
Entomology 14:705-708.
In Kabale District, southwest Uganda, four sets of options for the integrated
control of bacterial wilt (BW) were tested in a participatory approach with farmers
between 1995 and 1999, along with experiments on crop rotation and soil
amendments. Options tested for integrated control were (1) an improved package
(IP) that consisted of clean seed, a less-susceptible variety (Victoria), and improved
cultural practices, and (2) a farmer package (FP), which consisted of a farmers'
variety and farmers' seed, planted under the farmer's cultural practices. Two
additional packages were later added: clean seed of Victoria planted under farmers' cultural practices, and farmers' variety and seed planted under improved
cultural practices. In the rotation trial, one-season rotation experiments were
conducted on mildly infested fields and two-season rotation experiments were
conducted on heavily infested fields. In the soil-amendment experiment, organic
materials from Sesbania sesban and Leucaena diversifo/ia were applied in amounts
sufficient to supply 100 kg N/ha either singly or combined with P and PK. Also
added were NP and NPK from inorganic sources at rates that would supply 100
kg/ha for each element. All three options for the integrated control of BW significantly reduced wilt and increased yield; With IP performing best. The options
were economically beneficial, with marginal rates of return of 1034% for IP,
805 % for clean seed of a less susceptible variety under farmers' practices, and
634% for improved cultural practices with farmers' variety and seed. A one-season
rotation in mildly infested fields significantly reduced wilt to 3.2-12.4 % , compared to 63 % in the control, and resulted in corresponding significant in creases in
total and marketable yields. Similarly, the two-season rotation in heavily infested
plots also resulted in significant reductions in wilt incidence and increases in yield,
with increases in marketable yields ranging from 216-400%. Planting two different crops in two consecutive seasons resulted in superior wilt control, compared
to planting the same crop consecutively. Applying organic materials did not
necessarily reduce BW incidence, but there was a noticeable effect when organic
materials were applied together with inorganic fertilizers. The best results in
relation to both disease control and increased yield were obtained when sesbania
was applied together with P and K.
The potato (Solanum tuberosum) is an
important food and cash crop in the
highlands of Uganda. lt is mainly grown in
1
129
the leading producer of the crop, accounting for over 42% of the national annual
production (Bariyanga, 1997), and in this
district potato ranks first as a cash crop
and fourth as a food crop (Lemaga et al.,
1999).
Although potato yields are as high as from
20 to 40 t/ha at the research station, the
national mean yield is about 7 t/ha (FAO,
2000), which is low compared with
production statistics in many countries.
The low yield is attributed to a number of
biotic and edaphic factors in addition to
weather and poor management. Bacteria!
wilt (BW), caused by Ralstonia
solanacearum, is the second most important yield-reducing biotic factor in Kabale
after late blight (Low, 1997; Lemaga et al.,
1999). Since BW cannot be controlled by
chemicals and persists in the soil for a
long time, it is increasingly becoming a
major threat to potato production in the
district, where it causes a yield loss of
about 26% (Lemaga et al., 1999) with
occasional loses of 100% if the disease
appears during the early stages of crop
growth (Opio, 1988; Kakuhenzire et al.,
1993). In Kabale, BW is caused by race 3,
biovar 2-A of R. solanacearum (data to be
published at a later date).
Race 3 has a narrow host range and can be
successfully controlled by integrated
disease management (IDM). The most
important components of IDM include
clean seed (Berrios and Rubirigi, 1993;
Van der Zaag, 1986), crop rotation (Yerma
and Shekhawat, 1991; Gunadi et al., 1998;
Lemaga, 2001 b), less-susceptible varieties
(French et al., 1997; Tusiime et al.,
1996a), and disease-free soil (Priou et al.,
1999a). BW can be eradicated if efficient
control measures are properly used, as
reported in Australia (Lloyd, 1976) and
Peru (French, 1994). The successful control
of BW in Kabale may, however, be more
difficult than we anticipate because of
small and very fragmented landholdings
that limit crop rotation and sanitation, and
because of poor soil fertility (Lemaga et
130
Research on Potato
The experiment was laid out in a randomized complete block design. Within a
season, each farmer's field (farm) was
considered as a replication; the number of
farmers per season averaged 14. For each
plot, the soil type and history of crop
rotation were recorded.
Economic analysis
Since adoption of any technology by
farmers is dependent on the economic
benefit they receive, analyses of partial
budget and marginal rates of return were
done for each of the 1DM options. Twentytwo farmers who participated in o-farm
integrated BW control experiments also
participated in this study. The costs of
131
The experiment was laid out in a randomized complete block design with three
replications. Plot size was 16.2 m2 , in
which disease-free Victoria patato seed
was planted and amended with either
organic materials from Sesbania sesban
(Sesbania) or Leucaena diversifolia
(leucaena) inorganic fertilizers (nitrogen
(N), phosphorus (P), potassium (K)), or
different combinations of these organic
and inorganic fertilizers (Table 1). To allow
time far decomposition, fresh sesbania and
leucaena leaves and twigs were incorporated into soil a week befare planting in
amounts needed to supply 100 kg N/ha.
lnorganic fertilizers N, P, and K were sidedressed at rates of 100 kg/ha each as urea,
triple super phosphate (TSP), and murate of
potash (KCI), respectively. Urea was split-
applied in equal halves at planting and
one month after planting, while TSP and
KCI were applied at planting. Late blight
was controlled with faur sprays of Dithane
M45 and one spray of Ridomil MZ, both at
recommended rates, beginning with the
appearance of the first disease symptoms.
The methods used to assess and col lect
data on BW incidence, yields, and statistical analyses were similar to those used in
the IDM experiment.
Crop rotation
132
Research on Potato
Lignin (%)
8.1
4.3
6.6
14.1
One-season rotation
Potato-on ion-potato
Potato-peas-potato
Potato-cabbage-potato
Potato-sweetpotato-potato
Pota to-mil let-potato
Potato-carrots-potato
Potato-beans-potato
Potato-potato-potato
Two-season rotation
Potato-beans-wheat-potato (P-B-W-P)
Potato-beans-maize-potato (P-B-M-P)
Potato-wheat-maize-potato (P-W-M-P)
Potato-beans-beans-potato (P-B-B-P)
Potato-maize-maize-potato (P-M-M-P)
Potato-wheat-wheat-potato (P-W-W-P)
Potato-potato-potato-potato (P-P-P-P)
Plots were laid out in a randomized
complete block design with three replications for the one-season experiment and
four replications for the two-season
experiment. In the two-season experiment,
either two different rotational crops or the
same crop were grown in two consecutive
seasons. Clean seed of the potato variety
Victoria was used for both trials, and local
varieties were used for the rotation crops.
All the crops were planted at recommended spacing, which for potatoes was
75 x 30 cm. Late blight was controlled
with four sprays of Dithane M45 and one
spray of Ridomil MZ, both at recommended rates, starting from appearance of
133
Table 2. Effects of the various integrated 8W control options on wilt incidence, total and marketable yields,
and percent increases in marketable yields during seasons 1996A, 19968, 1997A, and 19978, Kabale,
Uganda.
Total
Marketable
lncrease in
Options
Wilt
incidence {%}
marketable yield {%}
yield {t/ha}
yield {t/ha}
1996A
IP
FP
IVCSFC
FVFSIC
LSDo.os
(2.3)
(3.7)
(3.2)
(3.4)
(NS)
17.10
12.10
14.60
10.10
3.20
16.50
12.30
14.40
10.40
4.10
15.80
27.80
20.73
24.10
(3.58)
(4.87)
(3.68)
(4.39)
(0.88)
15.80
10.80
11.70
13.50
2.34
14.80
9.90
10.80
12.10
2.25
49.50
6.05
20.73
9.98
17.12
(1.82)
(3.80)
(2.49)
(3.49)
(0.70)
10.35
7.56
7.87
9.67
1.94
10.13
7.28
7.70
9.27
2.00
39.20
3.26
21.09
3.73
10.94
(1.55)
(3.79)
(1.61)
(2.88)
(1.70)
6.10
4.90
4.94
6.40
ns
5.78
3.80
4.70
5.40
ns
5.30
13.40
9.70
11.00
34.10
17.10
-15.40
19968
IP
FP
IVCSFC
FVFSIC
LSDo.os
9.20
22.00
1997A
IP
FP
IVCSFC
FVFSIC
LSDo.os
5.77
27.30
19978
IP
FP
IVCSFC
FVFSIC
LSDo 05
52.10
23.70
42.10
Notes: Figures in parentheses are square roots of wilt incidence. NS= not significant. IP = improved package,
FP = farmer package, IVCSFC = improved variety and clean seed planted under farmers' cultural practices, FVFSIP =
farmers' variety and seed planted under improved cultural practices.
134
Research on Potato
Table 3. Analysis of partial budget and marginal rates of return far traditional and improved technology
options.
Components
.lntegrated BW control options
IP
FP
IVCSFC
FVFSIC
Yield {t/ha)
13.92
9.37
11.34
11. 70
Amount sold as seed {t/ha)
9.48
4.22
7.34
4.50
Amount soldas ware {t/ha)
4.45
5.15
4.00
7.20
lncome from seed (US$/ha)
3385.71
1205.71
2621.43
1285.71
lncome from ware (US$/ha)
847.62
980.95
781.90
1371.43
Gross income (US$/ha)
4233.33
2186.67
3382.33
2657.14
Costs that vary (US$/ha)
884.46
69.43
836.13
768.03
Net benefit (US$/ha)
3346.87
1482.68
2547.21
1889.11
Marginal rates of return (%)
1034.00
805.60
634.50
Note: IP = improved package, FP = farmer package, IVCSFC = improved variety and clean seed planted under farmers'
cultural practices, FVFSIP = farmers' variety and seed planted under improved cultural practices.
Soil amendments
The incidence of bacteria! wilt varied with
the seasons and was highest in 1998A and
lowest in 1999A (Table 4), differences that
could be attributed to variations in rainfall
since wilt incidence is positively correlated to precipitation (Akiew, 1986). The
control plots were the worst affected,
Crop rotation
One-season rotation. In the mildly infested
field (15-20% infestation) all the oneseason rotations with pulses, cereals,
vegetables, and root crops, although not
135
Table 4. Effect of soil amendments on patato tuber yield and incidence of bacteria! wilt during the 1998A,
1998B, and 1999A seasons, Kabale, Uganda.
Marketable
% Relative increase
Total tuber
Wilt incidence
Treatment
in marketable yield
yield (t/ha)
(%)
t/ha)
1998A
L
S+PK
L+PK
S+P
L+P
NPK
NP
62.60
62.00
49.90
51.40
48.20
51.60
44.70
31.60
66.50
(7.9)
(7.9)
(7;0)
(7.1)
(6.9)
(7.2)
(6.7)
(5.6)
(8.2)
(0.9)
9.9
8.2
13.7
10.5
9.6
7.3
9.0.
11.2
4.9
5.1
8.0
6.1
10.9
7.6
7.6
4.7
7.7
9.9
3.3
4.8
142.4
84.5
230.3
130.3
130.3
42.4
133.4
200.0
14.44
21.21
4.47
19.80
17.46
22.44
16.71
12.17
24.48
(3.8)
(4.6)
(2.1)
(4.4)
(4.2)
(4.7)
(4.1)
(3.4)
(4.9)
(0.5)
25.9
23.1
30.9
26.4
23.9
22.3
27.8
19.9
11.5
3.8
24.2
20.4
29.6
23.2
20.8
19.3
25.2
18.3
9.3
4.8
160.2
119.4
218.3
149.5
123.6
107.5
171.0
96.8
6.30
7.80
2.90
7.40
12.20
7.35
7.03
7.47
5.30
(2.5)
(2.8)
(1.7)
(2.7)
(3.5)
(2.7)
(2.6)
(2.7)
(2.3)
NS
18.3
19.1
22.5
19.5
17.9
16.9
19.5
18.7
17.4
3.6
17.7
18.3
21.9
18.5
17.2
16.0
18.4
17.4
16.5
3.8
7.2
10.9
32.7
12.1
4.2
-3.0
11.2
5.4
LSD(aUi)
19988
L
S+PK
L+PK
S+P
S+P
NPK
NP
LSD(QUi)
1999A
L
S+PK
L+PK
S+P
L+P
NPK
NP
LSD(o.(
Note: Figures in parentheses are square roots of wilt incidence. NS= not significant at P < 0.05. S = Sesbania sesban,
L = Leucaena divesifolia, N = nitrogen, P = phosphorus, K = potassium, C = control. (Reprinted with permission of
the African Crop Science Journal from Lemaga et al. (2001 a).)
136
Research on Potato
35
cu
.!::
25
"O
Q)
::;., 20
Q)
15
<ti
15
()
~
30
10
Y= -2.5281x + 26.97
R2 = 0.56
(P<0.001)
#t
f4~
10
Table 5. The effect of a one-season (1999A) rotation with various crops on incidence of bacteria! wilt, tuber
yields, and increase in marketable tuber yields, Kachwekano, Kabale, Uganda.
lncrease in
Treatment
Bacteria!
Total yield
Marketable
marketable yield (%)
wilt (%)
(Vha)
yield (Vha)
80
Potato-onions-potato
12.4 (3.5b)
20.4a
20.0a
67
Patato-peas-patato
5.0 (2.2b)
18.6a
18.5a
59
Potato-cabbage-potato
6.9 (2.6b)
17.7a
17.6a
48
Potato-sweetpotato-potato
3.8 (1.6)
16.4ab
16.4a
50
Potato-millet-potato
3.2 (1.7b)
16.?ab
16.?a
84
Potato-carrots-potato
11.0 (3.3b)
20.4a
20.4a
63
Potato-beans-potato
7.4 (2.2b)
18.1a
18.1a
Potato-potato-potato
62.2 (7.8a)
12.4b
11.1b
Note: Figures in parentheses are square roots of wilt incidence. Means followed by the same letter in columns are not
significantly different at 0.05 probability level. Adapted from Lemaga et al (2001b).
137
90
80
70
l
-~
cu
60
50
40
13
ctl
co 30
20
10
o
40
47
54
61
68
75
82
89
96
Conclusions
The studies reported here demonstrate that
bacteria! wilt can be satisfactorily con-
Table 6. Effect of two-season rotation with cereals and beans on incidence of bacteria! wilt and patato yields
at Kachwekano, Kabale.
Wild incidence
Total yield
Marketable
Treatment
lncrease in
(t/ha)
yield (t/ha)
marketable yield (%)
Potato-beans-wheat-potato
36.4 (5.81 bcd)
9.27a
7.40a
300
Potato-beans-maize-potato
21.9 (4.52d)
10.37a
9.00a
386
Potato-wheat-maize-potato
25.2 (4.89cd)
10.?a
8.90a
381
Potato-beans-beans-potato
40.3 (6.32bc)
8.92a
6.68a
216
Potato-maize-maize-potato
49.4 (7.01b)
10.15a
8.40a
354
Potato-wheat-wheat-potato
40.0 (6.28bc)
10.73a
9.23a
400
Potato-potato-potato-potato
81.1 (8.96a)
3.20b
1.85b
Note: Figures in parentheses are square roots of wilt incidence. Means followed by the same letter in columns are not
significantly different at 0.05 probability level by Duncan's Multiple Range Test.
Adapted from Lemaga et al. (2001 b).
138
Research on Potato
Acknowledgements
This study was conducted under the
African Highlands lnitiative, which also
provided the funds. The excellent collaboration of the National Agricultura!
Organisation of Uganda is gratefully
acknowledged. The valuable technical
support of Drs. S. Priou, E.R. French,
P. Ewell, M. Olayna, R. El-Bedewy, and
J.J Hakiza, both during the execution of
the experiments and in the preparation of
this manuscript, is highly appreciated.
The author is also indebted to Mr. Geofrey
Manzi for data collection and Mr.
R. Kakuhenzire and Patrick Okori for their
support in data analysis.
References
Akiew, E.B. 1986. lnfluence of soil
moisture and temperature on the
presence of Pseudomonas
solanacearum. In: Persley, G.J. (ed.).
Bacteria! wilt disease in Asia and the
South Pacific. ACIAR Proceedings No.
13. Australian Centre for lnternational
Agricultura! Research, Canberra,
Australia. p. 77-79.
Bang, S.K. and G.C. Wiles. 1995. Control
of bacteria! wilt (Pseudomonas
solanacearum) in potato by crop
rotation. In: Rasco, E.T. Jr. and
F.B. Aromin (eds.). SAPRAD on the third
year of phase 111. Selected research
papers. Vol. 1. South Asian Program for
Potato Research, Manila, Philippines.
p. 103-106.
Berrios, O.E. and A. Rubirigi. 1993.
lntegrated control of bacteria! wilt in
seed production by the Burundi national
potato program. In: Hartman, G.L. and
A.C. Hayward (eds.). Bacteria! wilt.
ACIAR Proceedings No. 45. Australian
Center for lnternational Agricultura!
139
140
Research on Potato
13 p.
Priou, S., L. Gutarra, H. Fernandez, and
P. Aley. 1 999a. Sensitive detection of
Ralstonia so/anacearum in latently
infected potato tubers and soil by
postenrichment ELISA. In: lmpact on a
changing world: CIP Program Report
1997-1998. lnternational Potato Center,
Lima, Peru. p. 111-122.
Priou, S., P. Aley, E. Chujoy, B. Lemaga,
and E. French. 1999b. lntegrated control
of bacteria! wilt of patato: CIP Slide
Training Series IV-3. lnternational Potato
Center, Lima, Peru. 30 p.
Ramesh, C.R. and A.K. Bandyopadhyay.
1993. Bacteria! wilt potential of
Andaman and Nicobar lslands. In:
Hartman, G.L. and A.C. Hayward (eds.).
Bacteria! wilt. Proceedings of the
lnternational Bacteria! Wilt Symposium
held at Kaohsiung, Taiwan, 28-31
October 1992. ACIAR Proceedings No.
45. Australian Centre for lnternational
141
143
144
Research on Patato
Sample testing
Tubers were randomly divided into 24
composite samples of 25 tubers each. For
the detection of R. solanacearum, tubers
were processed as described by Priou et al.
(1999a, 1999b) in the laboratory of the
Peruvian Crop Protection Service
(SENASA) in Cajamarca, Peru, and in the
laboratory of the Foundation for the
Promotion of and Research on Andean
Products (PROINPA) in Sucre, Bolivia.
After tuber disinfection, fragments were
taken from the vascular ring of each tuber;
the fragments from a composite sample of
25 tubers were crushed together in one
bag. Two 0.5 mi aliquots of tuber extract
were taken from each bag and rn ixed with
an equal volume of modified SMSA broth
for enrichment (incubation for 48 h at 30C
with manual agitation at least twice a
145
Table 1. Detection of Ralstonia solanacearum in seed lots coming from fields with various bacteria! wilt
(BW) incidences in Cajamarca and La Libertad departments, Peru.
Variety
ELISA-Positive among
Si te
Altitude {m)
Lot no. Bacteria! Wilt in
24 composite samples
the field {%)
o
Canchn
La Encaada
2800
5
1
o
Canchn
La Encaada
3000
7
2
Huamachuco 1
o
Canchn
3000
8
3
Amarilis
San Juan
2200
o
4
5
Cajamarca
o
3
Amarilis
2750
5
o
Canchn
La Encaada
3000
14
6
o
Amarilis
Huamachuco 1
3000
7
6
o
o
Amarilis
Contumaza
2700
8
Yungay
o
o
La Encaada
3000
9
o
10
Perricholi
Parean cooperative
3150
o
o
11
4
Tumbay
Chota
2900
o
1
Atahualpa San Miguel
12
2800
13
o
o
Huagalina
Su ere
2650
o
Huagalina
Su ere
2900
o
14
o
o
15
Canchn
Huamachuco 1
3200
Huamachuco 1
o
o
Canchn
16
3350
o
Yungay
Huamachuco 1
17
o
3400
o
18
Amarilis
Huamachuco 1
o
3200
Huamachuco 1
19
o
o
Yungay
3150
o
Huamachuco 1
20
Canchn
o
3200
o
21
Canchn
o
Huamachuco 1
3200
o
Yungay
22
8
La Encaada
2900
23
o
Yungay
11
Baos del Inca
3000
o
24
4
Amarilis
La Encaada
3000
o
25
6
Amarilis
La Encaada
3100
o
26
o
Amarilis
Baos del Inca
2800
27
o
o
Amarilis
Baos del Inca
2900
o
Yungay
28
13
Baos del 1nea
2950
o
29
Amarilis
10
La Encaada
3100
o
30
7
Amarilis
La Encaada
3000
o
o
31
Amarilis
Su ere
2650
o
32
9
Amarilis
Su ere
2700
o
Canchn
Sucre
o
33
2900
o
Yungay
34
Baos del Inca
5
3000
Total
126
0.1-1
1
35
Amarilis
Llaca nora
2900
0.1-1
36
15
Amarilis
Su ere
2850
37
0.1-1
19
Canchn
Su ere
2600
0.1-1
18
Canchn
38
Eduardo Villanueva
2200
39
0.1-1
17
Canchn
Sucre
2750
40
0.1-1
3
Canchn
Eduardo Villanueva
2240
0.1-1
Canchn
41
8
Sucre
2650
42
0.1-1
8
Mixture
Su ere
2700
43
0.1-1
Chaucha
5
San Pablo
2700
44
1.1-3%
11
Canchn
Jos Galvez
2600
Canchn
45
1.1-3%
13
Su ere
2650
46
1.1-3%
18
Mixture
Su ere
2650
47
1.1-3%
13
Mixture
Su ere
2700
48
1.1-3%
21
Amarilis
Sucre
2600
49
1.1-3%
3
Amarms
San Juan
2200
1.1-3%
15
50
Amarilis
Llaca nora
2900
Total
188
1
146
Research on Potato
Calculations
For a given bacteria! wilt incidence (%),
the average number of positive composite
(25-tuber) samples per lot, M, could be
calculated as M = number of positive
composite samples/number of positive lots.
This average gives us M = 24 p where p is
the probabi 1ity that the composite sample
in a lot is positive. Thus p = M/24 and
from p we obtain q = 1 - p (probability of
obtaining no infected sample). Then
applying the binomial distribution, the
probability P to obtain at least one infected composite sample (that is enough to
consider the lot infected) is P = 1 - qx
where X (in this case 24) is the number of
25-tuber samples to be analyzed. Different
values of X are tested and the optimum
sample size is the number X that al lows a
probability (P) of detection of 95%
(Snedecor and Cochran, 1980).
Results
Both in Peru and Bolivia, R. solanacearum
was detected in several composite
samples of all lots that carne from fields
with visible symptoms, even from those
with very low levels of wilt incidence
(Tables 1 and 3). For the lots coming from
fields with O. 1-1% wilt incidence, a
probability of 95% to detect one positive
sample was reached by analyzing 150
tubers in Peru and 225 tubers in Bolivia
(Tables 2 and 4). For fields with between
1. 1 and 3% wilt, 100 tubers have to be
analyzed in Peru and 200 in Bolivia to
obtain P = 95% (Tables 2 and 4).
In Peru, 18 seed lots (53.3%) coming from
apparently healthy fields were found
positive in ELISA (Table 1). Five of these
positive lots were from recognized seed
growers. Lots 8 and 1O, which served as
tentative negative controls, were found
negative. All positive results in ELISA
were confirmed by isolation of the pathogen in Cajamarca or by PCR at CIP;
pathogenicity tests on tomato were also
positive (data not shown).
In Bolivia, 15 seed lots (35.7%) of the
symptomless crops were found infected
with BW (Table 3). From the 42 apparently
healthy lots, 28 were from highlands
(3284-3450 m) and 14 were collected
between 2371 and 2850 m. Among the 14
lots from lower areas, 50% were found
positive. This is not surprising because BW
is endemic in the seed-producing areas of
147
Table 2. Probability (P) of detecting Ralstonia so/anacearum in seed lots coming from fields with various
BW incidences according to the sample size analyzed in Peru.
P for 0.1-1% ewz P for 1.1-3% ew3
Composite samples {X)
Tubers
P for 0% ew1
2200-3350 m
2200-2900 m
2200-2900 m
25
0.292
1
0.435
0.560
50
75
3
4
5
6
7
8
9
100
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
1P =
2p =
1 - o. 708x.
1 - 0.56sX.
3p=
1 - 0.440X.
125
150
175
200
225
250
275
300
325
350
375
400
425
450
475
500
525
550
575
600
0.499
0.645
0.749
0.822
0.874
0.911
0.937
0.955
0.968
0.978
0.984
0.989
0.992
0.994
0.996
0.997
0.998
0.999
0.999
0.999
0.999
1.000
1.000
Cordillera del Rosal, Chuquisaca Department. From the 28 lots from the highlands
in Chuquisaca and Potosi, 8 (28.6%) have
been found positive, all from Chuquisaca
(Table 3). This result demonstrates that
infected seed coming from the contaminated inter-Andean valleys of Chuquisaca
is responsible for contamination of seedproducing areas. The fact that lots from the
highlands of Potosi were all found negative was expected because BW has never
been reported in this area. These negative
results confirm the specificity of the test
(absence of cross-reaction in ELISA).
In Peru, the analysis of 225 tubers gives a
probability of 95% to detect at least one
positive sample in lots harvested from
symptomless fields (Table 2). When data
were separately analyzed by year (14 lots
analyzed in 1999 and 20 lots in 2000),
148
Research on Potato
0.681
0.820
0.898
0.942
0.967
0.982
0.990
0.994
0.997
0.998
0.999
0.999
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
0:806
0.915
0.963
0.984
0.993
0.997
0.999
0.999
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
Discussion
The serological detection technique. has
been val idated because the pathogen
could be detected in all fields with visible
BW symptoms. No .cross-reaction was
observed at Cajamarca (all positive RISA
results were confirmed), anc:Fexpected
negative seed lots were found negative in
both countries.
Table 3. Detection of Ralstona solanacearum in seed lots coming from fields wtth various BW incidences in
seed-producing areas of Chuquisaca and Potosi departments, Bolivia.
lot no.
Bacteria! Wilt in
ELISA-positive among 24
Site
Altitude (m)
the field (%)
composite samples
o
o
1
Canadillas
2850
o
o
Canadillas
2
2850
o
o
Canadillas
2850
3
o
o
Canadillas
4
2850
o
Lajas
2
3284
5
o
o
Huano loma
3284
6
o
Lajas
o
3284
7
Lajas
o
3284
1
8
o
o
Tarabuco
3284
9
3284
1
Tarabuco
o
10
Alto leuquepa.
o
2600
o
11
Alto Rosal
2500
o
o
12
Alto leuquepa.
2600
o
o
13
Alto Rosal
2500
1
o
14
2371
Faje ha
1
o
15
Fajcha
2390
o
3
16
2490
Alto Rosal
3
o
17
2390
Fajcha
2
o
18
2371
Faje ha
1
o
19
2385
Faje ha
1
o
20
3284
Tarabuco
o
o
21
3284
Tarabuco
1
o
22
3284
Lajas sijlla
o
o
23
3284
Lajas
o
o
24
3284
Lajas
o
o
25
3284
Lajas
o
o
26
3284
Lajas
o
o
27
3284
Lajas
o
o
28
3284
Jatun Mayu
2
o
29
3400
Central Alta
o
o
30
3400
Maraza
o
o
31
3400
Ckochas 1
o
o
32
3400
Simi simi 1
o
o
33
3400
Sijllani 1
o
o
34
3400
Chacabuco 1
o
o
35
3450
Chinoli 1
o
o
36
PROINPA-E.E.Chinoli1
3450
o
o
37
3400
Chacabuco 1
o
o
38
3400
Kawawasi 1
o
o
39
3450
Lajas (Tarabuco)
2
o
40
3450
Lajas (Tarabuco)
1
o
41
3450
Lajas (Tarabuco)
1
o
42
23
Total
2400
Muyuorko
5
0.1-1
43
1875
Las Palmas
16
0.1-1
44
2400
Muyuorko
10
0.1-1
45
2315
Muyuorko
6
0.1-1
46
2410
Muyuorko
3
0.1-1
47
2400"
Muyuorko
8
0.1-1
48
2450
Chajra Mayu
8
1.1-3
49
149
Table 3. (continued)
Lot no.
Bacteria! Wilt in
the field (%)
ELISA-positive among 24
composite samples
1.1-3
1.1-3
1.1-3
1.1-3
1.1-3
1.1-3
1.1-3
5
3
9
6
8
9
7
103
50
51
52
53
54
55
56
Total
1
Site
Muyuorko
Muyuorko
Chajra Mayu
Muyuorko
Muyuorko
K'ara Estancia
Las Palmas
Altitude (m)
2390
2400
2450
2390
2395
2468
2200
Table 4. Probability (P) of detecting Ralstonia solanacearum in seed lots coming from fields with various
BW incidences according to the sample size analyzed in Bolivia.
P for 0.1-1% ewz P for 1.1-3% ew3
Composite
Tubers
P for 0% ew1
2200-2468 m
samples (X)
2371-3450 m
1875-2410 m
0.307
0.287
1
25
0.060
0.520
50
0.116
0.492
2
0.667
3
0.638
75
0.169
4
100
0.219
0.769
0.742
0.840
0.816
5
125
0.266
0.889
0.869
150
0.310
6
0.923
0.906
7
175
0.352
8
200
0.390
0.933
0.947
9
225
0.427
0.952
0.963
10
250
0.461
0.966
0.974
11
275
0.494
0.976
0.982
12
300
0.524
0.983
0.988
13
325
0.553
0.988
0.991
14
350
0.579
0.991
0.994
15
375
0.605
0.996
0.994
16
400
0.628
0.996
0.997
0.998
0.997
17
425
0.651
18
450
0.672
0.999
0.998
19
475
0.691
0.999
0.998
0.999
0.999
20
500
0.710
1.000
0.999
21
525
0.727
1.000
0.999
22
550
0.744
1.000
1.000
23
575
0.759
1.000
1.000
24
600
0.773
1 P = 1 - 0.94QX.
2 p = 1 - 0.66flX.
3 P = 1 - 0.713x.
150
Research on Potato
Conclusion
Sample sizes of 250 and 350 tubers have
been used successfully to detect R.
solanacearum in Peruvian seed lots from
symptomless potato crops with probabilities of 95% and 99%, respectively. These
are numbers of seed that feasibly can be
processed in a seed-health test without
incurring too high a cost of labor and
material.
The number 400 has been selected by
many seed certification agencies. According to our results in Peru (Table 2), the
European standard of 200 tubers would
provide a PEA of 6.3% (P = 0.937) in the
case of symptomless crops. That is slightly
h igher than the theoretical probabi 1ity of
5%. Considering that moderately to highly
susceptible potato varieties can harbor
latent infection in their progeny tubers at
frequencies ranging from 20% to 70%
(Priou et al., 2001 ), field incidences of 1 %
and 0.1 % wou Id lead to percentages of
infected tubers of 0.2-0.7% and 0.020.07%, respectively. According to the
binomial distribution, the number of tubers
theoretically needed for testing to detect
these levels of tuber infection at the 95%
level of confidence would be much higher
than that necessary in Peru to detect the
same tuber infection rates (Table 5).
Table 5. Number of tubers (n) to be analyzed to detect Ralstonia solanacearum in seed lots with various
tuber infection levels (p) at confidence levels P of 95% and 99%.
n theoretical 1
n observed
p' = wilt
p = infected
1-p
P
incidence
tubers rate
1360
100
0.9978
0.95
0.011
0.0022
100
0.95
388
0.9923
0.011
0.0077
14977
150
0.95
0.9998
0.001
0.0002
150
4278
0.95
0.9993
0.001
0.0007
0.011
0.011
0.001
0.0022
0.0077
0.0002
0.9978
0.9923
0.9998
0.99
0.99
0.99
0.001
0.0007
0.9993
0.99
2091
596
23024
6577
150
150
200
200
1 n theoretical was calculated following the binomial distribution (P = 1 - [1-p]n) and compared to the sample sizes
(n observed) that were effective to detect these levels of infection in Peru, according to Table 2.
151
Research on Potato
References
Certification committee, The Patato
Association of America. 1985. Problems
in uniform seed patato certification
procedu'res. American Patato Journal
62:376-386.
Ciampi, L., L. Sequeira, and E.R. French.
1980. Latent infection of potato tubers
by Pseudomonas solanacearum.
American Patato Journal 57:377-386.
Clayton, M.K. and S.A. Slack. 1988.
Sample size determination in zero
tolerance circumstances and the
implications of stepwise sampling:
Bacteria! ring rot as a special case.
American Patato Journal 65:711-723.
De Boer, S.H. 1991. Current status and
future prospects of bacteria! ring rot
testing. American Patato Journal
68:107-113.
De Boer, S.H. and J.W. Hall. 1996. The
probability of detecting Clavibacter
michiganensis subsp. sepedonicus by
indexing seed patato lots with
serological tests. Journal of
Phytopathology 144:459-463.
French, E.R., L. Gutarra, P. Aley, and
J. Elphinstone. 1995. Culture media for
Pseudomonas solanacearum: lsolation,
identification and maintenance.
Fitopatologia 30:126-130.
153
155
Methodology
Several steps were necessary to carry out
this research: (1) samples were collected
and tested for thei r response to temperature and soil moisture, (2) a massmultiplication process was developed, and
(3) the mass-produced fungus medium was
field-tested in rustic potato stores.
Sampling
Three test sites were selected in the
Peruvian Andes: Chinchero, Cusco, in the
south; Aymara, Huancayo, in central Peru;
and La Encaada, Cajamarca, in the north.
For each site, 1O sample units were
established in fields previously planted to
potato and 10 more were established in
rustic potato stores. A sampling unit
consisted of one square meter of soi 1
surface. For sampling purposes, three
layers, 1O cm thick, were studied to
determine the depth at which the weevil
and the fungus occurred. The soil of each
subsample was sieved and the number of
Andean patato weevils (larvae, pupae, and
adults) was recorded. lt was also noted
whether the i nsects were healthy or
infected with B. brongniartii. Healthy
insects were taken to the laboratory and
stored under moist conditions for eight
days to verify their uninfected condition.
lnfected insects were also taken to the
156
Research on Patato
Results
Geographical distribution
B. brongniartii infection was found in
specimens of the Andean potato weevil in
all sampling units of field and rustic potato
stores in Cusca, Huancayo, and
Cajamarca. In the field, the average level
of infection found in the weevil population
ranged from 9% in Huancayo to 12.4% in
Cusco. In rustic stores, the level of infection ranged from 11 .4% and Huancayo
15.2% in Cajamarca to 27.1 % in Cusco
(Table 1).
157
120
100
Soil moisture 1O %
Temperature
f;
-~
>.
Q)
50
.o
158
Research on Potato
1 day
15 days
30 days
45 days
Discussion
B. brongniartii is a widely distributed
fungus in the Peruvian Andes, infecting
larvae and other 1ife stages of the Andean
patato weevil, Premnotrypes spp. This
study found that the larval stage of the
Andean patato weevil is the life stage
most commonly infected by the fungus,
confirming earlier observations (Rojas,
1981; Vera, 1992). Tests also confirmed
that the larval is the stage most susceptible to the fungus, which produces more
conidia when grown in the larvae than in
other developmental stages of the weevil.
Most weevils sampled during the larval
stage were found in the upper 10 cm of
soil, as previously reported (Alczar, 1976;
Muoz, 1998). lt was also found that
B. brongniartii can remain viable for more
than one year under cool, dry conditions in
infected larvae filled with conidia. After a
year, the fungus activated mycelial growth
when moisture was provided.
Temperature and soil moisture have an
influence on the onset of the disease and
the development of the fungus, enhancing
either mycelial growth (moist conditions)
or conidial production (dry conditions).
Th is i nformation provides a basis for the
mass production, use, and conservation of
References
Alczar, S.J. 1976. Biologa y
comportamiento del "gorgojo de los
Andes" Premnotrypes suturicallus
Kuschel (Coleop: Curculionidae).
Master's thesis. Universidad Nacional
del Centro del Per, Huancayo, Peru.
90 p.
Alczar, J., K.V. Raman, H. Torres, and
E. Yabar. 1 990. Beauveria spp. hongo
amigo del agricultor. Rev. Medio
Ambiente (Lima, Per) 45:44-46.
Alves, S. 1998. Controle microbiano de
insetos. Second edition. Funda<;ao de
Estudos Agrrios "Luiz de Queiroz"
(FEALQ), Piracicaba, Brazil. 1163 p.
Brady, B. 1979. Descriptions of pathogenic
fung and bacteria Beauveria
brongniartii and Beauveria bassiana.
Commonwealth Mycological lnstitute,
Surrey, UK. p. 602-603.
De Hoog, G.S. 1972. The genera
Beauveria/ Jsaria, Tritirachium. Study in
micology. Number 1. Kent, UK. 41 p.
Faull, J.L. 1986. Fung and their role in
crop protection. In: Biotechnology and
Crop lmprovement and Protection.
University of London, UK. p. 141-149.
159
160
Research on Potato
161
Results
Greenhouse tests
Research on Patato
Laboratory tests
In general, the effects recorded far all
treatments in the laboratory followed the
same trend as those in the greenhouse
(Table 2, Figure 1 ). That is, egg and larval
mortal ity decreased as the dosage of
abamectin alone decreased. The addition
of plant oil increased the mortality far all
abamectin dosages, with a clear trend of a
higher rate of increased mortality far lower
dosages of abamectin (Table 2). Comparing treatment results obtained in the
laboratory and in the greenhouse, sorne
quantitative differences can be noted. In
general, mortality of eggs and larvae was
higher for each treatment in the laboratory
than in the greenhouse. The ratio of
toxicity to the amount of oil added was
Table 1. Effeet of abameetin alone and abameetin-oil mixtures on eggs and larvae of the leafminer fly,
Liriomyza huidobrensis in greenhouse test.
Mortality (%)
Dosage (%)
Egg
Larvae 11
Larvae
Oil
Larvae 1
Abamectin
b,
o
66.0
d
81.3
ed
57.2
T1
0.15
79.6
o
77.3
e
69.8
de
65.6
T2
0.1125
58.1
e
58.1
e
24.3
0.075
o
42.6
d
40.9
e
T3
f
21.0
f
10.3
o
23.8
27.6
T4
0.0375
e
97.1
a
85.7
94.0
a
97.6
a
T5
0.15
1
91.4
94.1
ab
89.5
89.3
b
T6
0.1125
1
a
87.0
be
76.5
70.6
ed
0.075
1
90.0
a
T7
79.2
72.7
67.5
ed
ed
75.2
b
T8
0.0375
1
g
fg
12.8
11.8
15.9
o
1
13.9
f
T9
g
g
h
7.1
5.2
5.8
o
o
4.8
T10
111
d
ed
e
fg
a
a
b
be
fg
g
1 Means within the same column, followed by a common letter do not differ significantly at P=0.05, according to
Table 2. Effeet of abameetin alone and abameetin-oil mixtures on eggs and larvae of the leafminer fly,
Liriomyza huidobrensis in laboratory test.
Dosage (%)
Mortality (%)
Abamectin
Oil
Egg
Larvae 1
Larvae 11
Larvae 111
T1
0.15
o
92.5
b,
100.0
a
100.0
a
100.0
a
T2
0.1125
O
87.5
b
96.6
b
100.0
a
100.0
a
T3
0.075
O
65.0
e
82.7
d
92.0
e
63.0
d
T4
0.0375
O
38.6
d
45.9
e
42.9
d
41.2
e
T5
0.15
1
100.0
a
100.0
a
100.0
a
100.0
a
T6
0.1125
1
100.0
a
100.0
a
97.8
b
100.0
a
T7
0.075
1
92.2
b
93.9
b
97.6
b
95.8
b
T8
0.0375
1
87.2
b
90.0
e
91.3
e
82.4
e
T9
O
1
O.O
e
23.9
f
7.9
e
16.3
f
no o
o
O.O
e
O.O
g
5.9
e
3.7
g
1 Means within the same column, followed by a common letter do not differ significantly at P-0.05, according to
163
120
Laboratory
100
Greenhouse
--...........
...,
80
ast::::
o
...
_
-----~
60
Q)
40
_J
---- Abamectin
........_ Abamectin +ol
20
0.15
0.1125
0.075
0.15
0.0375
0.1125
0.075
0.0375
Dosages
Figure 1. Effect of abamectin and abamectin-oil mixtures on leafminer fly larvae, Liriomyza huidobrensis, under
laboratory and greenhouse conditions, Lima, Peru, 2000.
higher in the greenhouse than in the
laboratory tests (Figure 1). Finally, oil
alone had no effect on eggs but showed
sorne mortality (8% to 24%) at the larval
stage.
Discussion
Abamectin alone, applied at the commercially recommended dosage, showed a
satisfactory level of control of the eggs
and larvae of the leafminer fly, confirming
reports of severa! authors (Ochoa and
Carballo, 1993; Buxton and McDonald,
1994; Sotomayor, 1998). Lower dosages
were less effective, indicating that the
recommended dosage is close to
abamectin's action threshold. Abamectin's
effect on eggs apparently occurs when the
embryo is full-grown-when the sickleshaped mandibles of the newly formed
larvae are visible through the chorion of
treated eggs-as reported by Schuster and
Everett (1983) and Sotomayor (1998). In
many cases, larvae died during the process
of hatching, befare any damage to the
mesophyll tissue could be detected.
Apparently, abamecti n has no effect on
164
Research on Potato
Conclusions
This research confirmed that when used on
bean plants, abamectin alone is effective
against the eggs and larvae of the
leafminer fly (L. huidobrensis) at the
commercially recommended dosages.
The addition of plant oil to abamectin
improved the potency of the insecticide,
also allowing the dosage to be reduced.
The cost of effectively treating 1 ha is
reduced by more than 60%-from $131 .30
for abamectin alone to $53.70 for the
mixture of abamectin and plant oilthereby making it available to small-scale
potato farmers with limited resources.
References
Abro, G.H., R.A. Dybas, A. Green, and
D.J. Wright. 1988. Toxicity of
avermectin Bl against a susceptible
laboratory strain and an insecticideresistant strain of Plutella xylostella
(Lepidoptera: Plutellidae). Journal of
Economic Entomology 81 :1575-1580.
Anderson, T.E., J.R. Babu, R.A. Dybas, and
H. Mehta. 1986. Avermectin Bl:
lngestion and contact toxicity against
Spodoptera eridania and Heliothis
virescens (Lepidoptera: Noctuidae) and
potentiation by oil and piperonyl
butoxide. Journal of Economic
Entomology 79:197-201.
Buxton, J.H. and O.C. McDonald. 1994.
Chemical control of the South American
leaf miner, Liriomyza huidobrensis. In:
Proceedings-Brighton Crop Protection
Scrobipalpuloides absoluta
(Lepidoptera: Gelechiidae). Pesq.
Agropecuaria Brasilia 30:313-318.
Guerrero H., E.O. 1999. Efecto de la
abemectina en mezcla con aceite
vegetal agrcola sobre la mosca
minadora Liriomyza huidobrensis
Blanchard (Diptera: Agromyzidae) en el
cultivo de papa. Thesis. Facultad de
Ciencias Agropecuarias, Alimentarias y
Pesqueras, Universidad Nacional Jos
Faustino Snchez Carrin, Huacho,
Peru. (unpublished)
Horowitz, A.R., Z. Mendelson, and
l. lshaaya. 1997. Effect of abamectin
mixed with mineral oil on the
sweetpotato wh itefly (Homoptera:
Aleyrodidae). Journal of Economic
Entomology 90:349-353.
Lasota, J.A. and R.A. Dybas. 1991.
Avermectins, a novel class of
compounds: lmplications for use in
arthropod pest control. Annual Review
of Entomology 36:91-117.
Leibee, G.L. 1988. Toxicity of abamectin
to Liriomyza trifo/ii (Burgess) (Diptera:
Agromyzidae). Journal of Economic
Entomology 81 :738-40
Mujica, N. and F. Cisneros. 1997.
Developing IPM components for
leafminer fly in the Caete Valley of
Peru. In: Program Report 1995-96.
165
166
Research on Potato
The in-trust potato collection conserved at CIP has been characterized with a
variety of molecular markers to reduce genetic redundancies and identify
subsets important for both conservation and crop improvement. We have
recently been concerned with developing a potato (Solanum spp.) genetic
identification (PGI) kit for curatorial purposes. More than 70 microsatellite
markers were tested on a genetically diverse sample of potato germplasm.
We have selected a subset of the 18 most informative microsatellites according to the quality of the DNA products they amplify, their genome location,
and the level of polymorphism they detect. These assays have shown that the
microsatellites selected are ubiquitous in all seven cultivated potato species.
The PGI kit has also been shown to be useful to differentiate potato varieties
and cultivars. We have further used this marker set to identify genotype
ambiguities in the ex situ germplasm collection. The PG 1 kit will be routinely
used to produce a true-to-type reference for each new entry in our
germplasm collection and to attempt to minimize human error in germplasm
management.
The potato collection held in trust at
the lnternational Potato Center (CIP) is
mainly composed of cultivated accessions
accounting for up to 70% of the 5,094
clonal and true seed entries (Table 1, after
Huamn et al., 1997). Clonal accessions
of cultivated potato are maintained in the
field at CIP Huancayo and in vitro at La
Molina, Peru. Subsets of the collection
maintained in vitro are introduced to the
field regularly not only to monitor reliability of conservation procedures but to
provide new morphological data or trait
evaluation useful to geneticists and
breeders. Therefore, the correct identity of
each accession maintained in vitro is
highly relevant to both curators and
breeders at CIP.
1
167
168
Research on Patato
Accessions (no.)
268
170
48
Genotyped (no.)
6
21
8
10
56
97
8
4
13
31
2644
144
48
11
3527
1567
o
5
o
4
73
5094
economical to exchange. SSR alleles or
genotypes can be registered in molecular
marker databases, or with regu latory
agencies, to permit the development of
cumulative infarmation on varietal
fingerprints. Such a tool is needed to
complement or replace current methods of
variety description, which rely on environmentally sensitive morphological
characteristics. Microsatellite technology
is also an appropriate technology far
developing country laboratories and
breeding programs, which often have
modest expertise in molecular biology
techniques.
However, microsatellites are not yet the
perfect genetic marker because they entail
both limitations and assumptions that
cannot always be met. Limitations include
their high discovery cost, estimated to be
around US$300/microsatellite, and their
relatively high species-specificity as
opposed to RAPO and AFLP (Westman and
Kresovich, 1998). Microsatel 1ite use also
involves several assumptions. The first of
these is that band sharing indicates allele
identity. Although this has been shown to
be true in cases where the bands have
been sequenced, we cannot rule out that
single-nucleotide substitutions are present
between two comigrating bands. Another
case of incorrect allelic identity of
comigrating bands is due to the forward. backward steps during DNA polymerase
'slippage' that is responsible for microsatell ite al lele generation (Kruglyak et al.,
1998). Microsatellite alleles can thereby
revert to an original identical size for
genotypes that are in reality distinct. The
other important assumption is that the
presence of a band corresponds to one
copy of an allele. lt may actually correspond to more copies of the same alleles
but we cannot ascertain this. The ampl ification of microsatellites cannot be made
quantitative because it would be impractical to have to optimize each for
quantitative PCR. Probably only a number
of microsatellites would be amenable to
quantification of allele dosage, but such
use would have to include a new assumption of no-allelic preference in PCR
amplification. Hence, it seems unworkable for developing a procedure that
would permit the detection of different
dosages.
Molecular fingerprints will likely become
an essential tool in the inter-institutional
exchange of elite materials to assure the
identity of materials in international
testing programs, whether or not they are
protected by plant breeders' rights. They
will also add critica! resolution to patato
pedigrees, allowing the tracing or monitoring of successful alleles and allele
combinations in breeding programs where
only the fittest survive. Such historical
information as allelic constitution of
patato pedigrees is presently not readily
available but is potentially very valuable
for identifying successful progenitors of
important characteristics in multitrait
selection programs. Finally, molecular
pedigrees are well suited to graphical
representation such as genomic displays
and overlays. This will likely facilitate
their use. A valuable potential application
of SSR fingerprints of gene bank holdings
is the abi 1ity to query new entries for
unique diversity, along with or befare their
acquisition into already large and expensive ex situ col lections.
169
Data analysis
The polymorphic index content for each
microsatellite on the germplasm sample
was calculated according to the Nei
statistic (1973): PIC=1-L(p 2 ), where pi is
the frequency of the ith allele detected.
170
Research on Patato
Results
Transferability of microsatellites between
cultivated potato species
An original group of 70 microsatellites was
tested using a sample of cu ltivated potato
accessions comprising at least four genotypes from each of the nine taxonomic
groups. Ali microsatellite primers amplified successful ly (Table 1). A first positive
indication that we were amplifying
homologous microsatellite loci with those
of S. tuberosum subsp. tuberosum carne
from the comparison of al lele phenotypes.
As a matter of fact, each microsatellite
displays a specific al lele phenotype on
denaturing polyacrylamide gels, such as
discrete bands, double or triple bands, or
stutter bands. These phenotypes were
clearly conserved among the nine groups
tested, supporting the analogous nature of
the microsatellite locus. Southern blot was
performed for 1 3 m icrosatel 1ites to confirm
sequence homology among amplification
products (data not shown). Both of these
analyses are concordant with the conservation of microsatellite loci across
cultivated potato species.
'l::t
'l::t
,...
N
co N" (W') co
" Lt)oen(W') N"Nco Lt)"(O(W') NooLt)
(W')
Lt')
Lt)
o Lt)
N
(W')
Lt)
Lt)
(W')
bp ,...
o o o o "
o o o o o o
Lt')
(W')
" " " " " " " " " "
240
~:~
----:
~
241
199
187
184
Table 2. Microsatellite set used to fingerprint 73 native patato cultivars including polymorphic index content
(PIC) per marker, and location on the patato genetic map.
Chromosome Annealing Genotypes Alleles PIC
temperature (no.) detected
("C)
(no.)
SCRl1
code
Repeat motif
STM1049
STM2030
STM1064
STM2022
STM1053
STM1058
STM3023
STM1031
STPAc58
STM0019
STM0031
STM2013
STM1104
STM1017
STM3012
STM1106
STM0037
STM0030
(ATA) 6
(CA)3 (TA) 5
(TA) 12 (TG) 4 GT (TG) 5
(CAA)J ... (CAA)J
(TA) 4 (ATC) 5
(ATT) 5
(GA) 9 ... (GA) 8 ... (GA) 4
(AT)13
(TA) 13
(AT)] (GT) 10 (AT) 4 (GT) 5 (GC) 4 (GT) 4
(AC) 5 ... (AC)J (GCAC) (AC) 2 (GCAC) 2
(TCTA) 6
(TCT) 5
(ATT) 5
(CT) 4 ... (CT) 8
(ATT)13
(TC) 5 (AC) 6 AA (ACh (AT) 4
Compound (GT/GC) (GT) 8
1
1
11
11
111
111
IV
V
V
VI
VII
VII
VIII
IX
IX
X
XI
XII
57
55
55
53
55
55
50
55
57
47
57
55
57
53
57
55
53
53
73
73
73
73
22
22
22
73
73
73
73
73
22
73
73
22
73
73
7
5
6
5
4
4
5
8
10
29
10
13
7
3
6
6
10
13
0.555
0.447
0.581
0.665
0.670
0.324
0.689
0.591
0.802
0.882
0.737
0.815
0.798
0.483
0.645
0.749
0.784
0.879
Allele
size
(bp)2
184-254
180-209
188-199
184-241
168-177
113-125
177-201
265-325
231-277
83-239
155-205
146-172
168-183
132-136
168-213
142-196
75-99
122-188
1 SCRI =
2 bp =
171
populations tested so far, and has consistently revealed higher ploidy levels when
more al le les than expected were observed.
172
Research on Potato
Discussion
Potato genetic resources represent a large
pool of valuable alleles for a wide range
of important traits. Samples of these
genetic resources are maintained on-farm
and ex-situ in germplasm repositories. Both
conservation strategies have advantages
and disadvantages, but share the difficulty
of maintaining genetic resources without
losing or mixing sorne samples over time.
Such events do occur no matter how strict
the management procedures. The possibility of detecting these and resolving
ambiguities often relies on combinations
of the following sources of information:
(1) the original data of the collector,
(2) herbarium specimens, (3) records of
morphological evaluations, or (4) samples
kept by a third party. Therefore, genetic
fingerprints based on DNA markers have
been proposed as excel lent true-to-type
references. Among all the DNA markers
available, microsatellites provide distinct
advantages for patato genetic identification due to their high genetic information
content, high reproducibility, and simplicity of use.
Seventy microsatellites were tested, which
led us to identify 1 8 that are proposed here
as a potato genetic identification kit.
These microsatellites were selected on the
basis of the quality of the amplified
product they produce, their correspondence to single loci, their PIC, and the
benefits of using at least one marker/
patato chromosome. Since cross-species
amplification may lead to false positive
resu lts, we ha ve tested these
Hualash (adg) 4x
1
1
Ccompis (adg) 4x
lmilla Blanca (adg) 4x
Sani lmilla (adg)4x
Camotilla (gon) 2x
--
Huanquita (stn)2x
Puca Huayro (cha) 3x
Huagalina (adg)4x
Yana Suli (adg) 4x
Yana lmilla (adg) 4x
Papa Chanca (tbr)4x
Guincho Negra (adg) 4x
0.60
0.65
0.70
0.75
0.85
0.80
Similarity Coefficient
0.90
0.95
1.00
Figure 2. Dendrogram of the 22 native patato cultivars produced by genotyping with 18 microsatellites. Ploidy is
indicated next to each name: adg =S. tuberosum subsp. andigena, stn =S. stenotomum, gon =S.
goniocalyx, phu =S. phureja, cha= S. x chauca, tbr =S. tuberosum subsp. tuberosum.
microsatellites in a small sample of the
eight cultivated species of patato. Allele
phenotype and DNA:DNA hybridization
revealed homology between the
microsatellite amplification product
between species. Therefare, we consider
that these loci are conserved among the
patato species tested.
This PGI kit will have many uses. We have
shown here two examples of its immediate
use. Twenty-two native patato cultivars
were characterized molecularly. Unique
DNA fingerprint were obtained far all but
two pairs of native patato cultivars that are
closely related. Cultivars derived from
diploid and triploid species farmed a
173
Acknowledgments
The authors are grateful to Z. Huamn,
W. Roca, R. Gomez, A. Panta, and
J. Toledo for providing sources of plant
material. We also thank severa! technicians who have contributed to the
optimization of these protocols. This work
was initially made possible through a grant
from the United Nations Oevelopment
Programme, Special Project Program, and
has been partially supported through a
grant from the Bundesministerium fr
Wirtschaftliche Zusammenarbeit und
Entwicklung (BMZ) and Oeutsche
Gesellschaft fr Technische
Zusammenarbeit (GTZ) (project no
96.7860.8-001 .00).
References
CIP (lnternational Patato Center). 1997.
Ghislain, M., O. Zhang, and
M.R. Herrera (eds.). 1997. Molecular
biology laboratory protocols. Plant
genotyping. Genetic Resources
Department training manual.
lnternational Potato Center, Lima, Peru.
Ghislain, M., O. Zhang, D. Fajardo,
Z. Huamn, and R. Hijmans. 1999.
Marker-assisted sampling of the
cu ltivated Andean potato 5olanum
phureja collection using RAPO markers.
Genetic Resources and Crop Evolution
46(6):547-555.
Huamn, Z., A. Golmirzaie, and
W. Amoros. 1997. The potato. In:
Fuccillo, O., L. Sears, and P. Stapleton
174
Research on Potato
p. 21-28.
Huamn, Z., R. Ortiz, O. Zhang, and
F. Rodrguez. 2000. lsozyme analysis of
entire and core collections of 5olanum
tuberosum subsp. andigena potato
cultivars. Crop Science 40:273-276.
Kruglyak, S., R.T. Ourret, M.O. Schug, and
C.F. Aquadro. 1998. Equilibrium
distributions of microsatellite repeat
length resulting from a balance between
slippage events and point mutations.
Proceedings of the National Academy
of Science 95:10,774-10,778.
Milbourne, O., R.C. Meyer, A.J. Collins,
L.D. Ramsay, C. Gebhardt, and
R. Waugh. 1998. lsolation, characterization and mapping of simple
sequence repeat loci in potato.
Molecular Genetics 259:233-245.Powell, W., G.C. Machray, and J. Provan.
1996. Polymorphism revealed by simple
sequence repeats. Trends in Plant
Science 1 :215-222.
Provan, J., W. Powell, and R. Waugh.
1996. Microsatellite analysis of
relationships within cultivated patato
(Solanum tuberosum). Theoretical
Applied Genetics 92:1,078-1,084.
Nei, M. 1973. Analysis of gene diversity in
subdivided populations. Proceedings of
the National Academy of Science USA
70:3,321-3,323.
Sneath, P.H.A. and R.R. Sokal. 1973.
Numerical taxonomy: The principies
and practice of numerical classification.
W.H. Freeman, San Francisco, CA,
USA.
Tautz, O. and M. Renz. 1984. Simple
sequences are ubiquitous repetitive
components of eukaryotic genomes.
Nucleic Acid Research 12:4, 127-4, 128.
Westman, A.L. and S. Kresovich. 1998.
The potential of cross-taxa simplesequence repeat (SSR) amplification
between Arabidopsis thaliana L. and
crop brassicas. Theoretical Appl ied
Genetics 96:272-281.
Twenty potato (Solanum tuberosum L.) accessions were retrieved from three
to five years (without subculturing) in vitro storage, and 18 from 1-year
cryopreservation. Using 23 morphological descriptors, all accessions were
compared with original clones to verify if they were true-to-type. Materials
from in vitro storage were compared with clones maintained in the field, and
those from cryopreservation with clones maintained in vitro. Comparison of
cryopreserved materials was made using three types of propagules (plantlets,
stem cuttings, and tubers). Additionally, accessions from in vitro storage were
compared with samples maintained in the field using 14 highly polymorphic
potato microsatellite markers. Six accessions (four from in vitro storage and
two from cryopreservation) showed differences in multiple morphological
characters, suggesting a case of misidentification. In the remaining accessions, differences were observed for 1O descriptors of flowering degree and
color expression. Fewer accessions showing differences (four) were found
when tubers were used as planting material. Simple sequence repeats (SSR)
analysis of 20 accessions, retrieved from in vitro storage and compared with
original clones, resulted in five accessions showing remarkable DNA differences, again revealing possible misidentifications. Four were found different
from the original clones by morphological and molecular analyses; and, one
accession was found different at the molecular level, even though it showed
no morphological differences.
175
for the world community. CIP has constantly sought to upgrade its conservation
methods to minimize genetic changes in
the clones conserved. Growth retardation
and reduced temperature are the two
factors successful ly used for in vitro patato
conservation, extending the culture
transfer period to 3-4 years (Golmirzaie
and Toledo, 1999). Cryopreservation holds
a great potential for the long-term conservation of patato genetic resources (Benson
et al., 1996; Golmirzaie et al., 1999;
Schafer-Menurh et al., 1996; Steponkus et
al., 1992). Plant material can be stored
indefinitely at ultra low temperature (e.g.,
in liquid nitrogen), minimizing risk of
genetic changes in cells because, in
theory, chemical and physiological
changes are prevented. We have been
testing this approach for the past five
years, taking advantage of an experimental collection including nine species with
over 300 accessions.
Maintenance of genetic integrity is an
important requirement in gene bank
management. Therefore, assessment of
genetic integrity in plant propagation has
been conducted using morphological
descriptors. More recently, the application
of molecular markers has provided additional insight into the detection and
measurement of genetic variation (Kumar
et al., 1999).
176
Research on Patato
Cryopreservation was carried out accordi ng to Golmirzaie and Panta (1997b). Forty
shoot tips per accession were thawed after
one year of storage in liquid nitrogen.
From them, three clonal lines (C1, C2, and
C3) were used for morphological comparisons with original clones maintained in
vitro. Three sets of planting material
consisting of (1) plantlets grown in Jiffy-7
42 mm peat pellets (Jiffy Products of
America, lnc. Batavia, IL, USA), (2) stem
cuttings rooted in Jiffy-7 42 mm peat
pel lets, and (3) tubers obtained from the
field plants propagated in vitro were used
to establish field plantings. Ten plants/
treatment (accession/conservation method/
planting material) were planted in the
field; three were used for the morphological evaluation. All comparisons were
made under the same environmental
conditions.
Molecular comparison
Twenty accessions from in vitro storage
were compared with original clones from
Results
Morphological analysis
A total of 38 accessions were evaluated
for morphological characters. The dendrogram obtained by an initial cluster
analysis revealed six cases of
misidentifications (four from in vitro
177
Table 1. Microsatellite markers used far molecular analysis of 20 patato accessions retrieved from in vitro
storage in the gene bank held in trust at CIP.
Fragment size range (bp)2
Chromosome
Ali eles
PIC 3
SCRl1 code
VI
93-217
16
0.875
STM0019
STGBSS-1
VIII
130-142
7
0.812
V
203-242
0.711
5
STPAc58
XII
136-191
0.792
STM0030
8
VIII
243-262
0.811
STM1016-1
8
VII
179-205
STM0031
6
0.727.
4
1
0.662
185-205
STM1049
VIII
224-243
STWAX-21
8
0.799
VIII
169-182
0.804
STM1104
7
IX
212-268
0.811
STM1052
8
169-213
STM3012
IX
0.664
6
IV
181-201
STM3023
4
0.692
X
131-163
STM1106
7
0.780
11
185-244
STM2022
4
0.588
1 SCRI = Scottish Crop Research lnstitute.
2 bp = base pairs.
3 PIC (Polymorphic index content) = 1 -(p2) where pi is the frequency of the ith allele detected.
178
Research on Patato
Microsatellite analysis
Twenty potato accessions were DNA
fingerprinted using 14 microsatellite
markers. These are located on 1O of the 1 2
chromosomes of potato. Hence, a large
portion of the pota to geno me ..cou Id be
assessed for genetic variation. We have
obtained a total of 92 alleles. The number
of al leles/locus ranged from four
Differences in 2
(2/2)
(3/3)
d4
d4, d2, d12
d2, d4, d19, d20
d2, d3, d4
d4, d12, d13
d12
(2/3)
d4
(2/2)
Discussion
Differences in morphological characters
between in vitro and cryopreserved clones
and control clones could be attributed to
inconsistency in the criteria used to record
qualitative data. For example, descriptor
stem color stage 2 (green with few pig-
either way when it was difficult to determine the score far intermediate
expression. Variability in morphological
descriptors could also occur dueto physiological age of plants, presence of
diseases, and environmental factors
(Kumar et al., 1999; Potter and Janes,
1991 ). Therefare, descriptors with higher
percent variation, (stem color, flowering
degree, distribution of secondary flesh
color (Table 4)) are not useful far detecting
genetic changes or describing clonal
identity. In this study, the morphological
comparisons using three types of
propagules showed that tubers should be
preferred as planting material rather than
stem cuttings or plantlets. The probability
of detecting slight morphological differences, due to physiological conditions
rather than genetic changes, seems higher
179
180
Research on Potato
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'-'NO
Cl:l <O en
O M
cu
+-'
en
.g:::: ~
e.a... ~
o- o
>
~""3'" ~
Ct)
>
C\J
Cl:l
.::: <(
1-
1'
-U >.
-
-1<
>
o
>
<O
......
C\J
,....._
Q)
~en
>
>
CX)
$: ~
en +-'
'
Q)
'-'=
e::-
~-E
-1<
.........
Q)
u
u
-1<
Q)
~o
l.{)
O)
.E ..
...... U)
a:
>
o
>
e::
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Ct)
"O
>
o
>
o
-1<
o
>
o
>
N-
Q)
O> LO
+-'
o ...-....e::
.e
+-'
e::..__.. e::
e~
2-M
C.
Q)
Q)
=r- ......
.:=a_~
--e::x:
~~(!J
Cl e::
.
-r-N
en
1
m en X
...
Q)
Cl:l
g, 8
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t-
181
Conclusions
Morphological comparisons between in
vitro storage material and the original
clones maintained in the field revealed
differences in 9 of 20 genotypes assayed.
Of the nine genotypes detected, four were
judged to be misidentifications. The
remaining five showed slight morphological differences attributable to pathological
or environmental factors.
Morphological comparisons between
material cryopreserved for one year and
the original clones maintained in vitro
revealed differences in six of 18 genotypes assayed. Of these, two were
misidentifications. The remaining four
showed small morphological differences,
again attributable to pathological or
environmental factors.
The data obtained from morphological and
SSR analysis of original potato clones,
maintained in the field gene bank vs those
from in vitro storage for up to five years,
support the occurrence of misidentifications of potato accessions in the
different repositories. Microsatel 1ite
markers have been found to be a powerful
tool to complement the results obtained by
morphological comparisons. Both morphological and molecular methods are in
agreement in detecting misidentified
accessions rather than genetic i nstabi 1ities
(Ghislain et al. 2001 ).
Acknowledgements
The authors thank Merideth Bonierbale for
her valuable comments on the manuscript.
References
Angel, F., V. Barney, J. Tohme, and
W. Roca. 1996. Stabi 1ity of cassava
plants at the DNA level after retrieval
from 10 years of in vitro storage.
Euphytica 90(3):307-313.
Benson, E., M. Wilkinson, A. Todd,
U. Okura, and J. Lyon. 1996.
Developmental competence and ploidy
stability in plants regenerated from
182
Research on Potato
p. 165-178.
Gupta, P. and R. Varshney. 1999.
Molecular markers for genetic fidelity
during micropropagation? Current
Science 76:1308-131 O.
Harding, K. 1994. Molecular stability of
the ribosomal RNA gene in Solanum
tuberosum plants recovered from slow
growth and cryoconservation. Euphytica
55:141-146.
Huamn, Z., J. Williams, W. Salhuana,
and L. Vincent. 1977. Descriptors for the
cu ltvated pota to and for the
maintenance and distribution of
germplasm collections. lnternational
Board for Plant Genetic Resources,
Rome, ltaly. 47 p.
Kumar, M., R. Barker, and B. Reed. 1999.
Morphological and molecular analysis
of genetic stability in micropropagated
Fragaria x ananassa cv. pocahontas. 1n
vitro Cellular and Development
Biology. Plant 35(3):254-258.
70:3321-3323.
Piola, F., R. Rohr, and P. Heizmann. 1999.
Rapid detection of genetic variation
within and among in vitro propagated
cedar (Cedrus libani Loudon) clones.
Plant Science 141 :159-163.
Potter, R. and M. Jones. 1991. Molecular
analysis of genetic stability. In: Dodds,
J. (ed.). In vitro methods for
conservation of plant genetic resources.
Chapman and Hall, London, UK.
p. 71-91.
Schafer-Menurh, A., E. Muller, and
G. Mix-Wagner. 1996. Cryopreservation:
An alternative from the long-term
storage of old potato varieties. Potato
Research 39:507-513.
Steponkus, P., R. Langis, and S. Fujikawa.
1992. Cryopreservation of plant tissues
by vitrification. Advances in LowTemperature Biology 1 :1-16.
183
184
Research on Potato
J. Espinoza, M. Bonierbale1
185
186
Research on Potato
Group 1. The largest group of test materials (100 clones) was chosen from CIP's
'pathogen tested list' (PTL) of elite clones
and varieties (> 100), and was tested at
San Ramon and La Molina during winter,
1989. These materials are highly selected
tetraploid clones with diverse production,
resistance, and utilization traits. They all
carry resistance or tolerance to at least
one important pest, disease, or stress
factor.
Group 2. Another group was a sample of
42 clones taken from a diploid phureja/
stenotomum population. This population
was selected in the USA, from Solanum
phureja and 5. stenotomum parents, for
heat tolerance, adaptation to long days,
tuber appearance and high dry matter
(Gautney and Haynes, 1983; Haynes and
Haynes, 1983). These materials were
tested only at La Malina during winter of
1989.
Group 3. A third group of approximately
80 clones was selected from CIP's broadbased breeding populations, also
tetraploid, which are variously adapted to
Results presented here on general evaluation of clones for their processing qualities
are overviews from several years of trials,
so designs are not presented in detail. In
general, for the evaluation of the clonal
material, trials followed randomized
complete block designs (RCBD) with two
or three replications of single 1O hill row
plots. Yields were expressed as the weight
of tubers on a whole plot basis (not
reported here).
Tests for processing quality (see below)
were usual ly conducted 1O days after
harvest, once the chemical composition of
the tubers had stabilized. However, in
arder to assess the i mpact of storage
conditions on processing qualities, additional postharvest treatments were added
to two trials. A group of 16 late blight
187
188
Research on Potato
Results
Evaluation of advanced clones and
varieties for processing characteristics
(Group 1)
Dry matter content
Dry matter content evaluations of CIP's
pathogen tested list (PTL) clones and
varieties suggested the presence of
considerable genetic variability for this
characteristic (Figure 1 ). The highest
values for dry matter content in La Molina,
of about 24-26%, were observed for the
following varieties: AKK69.1 (720052),
Tomasa Condemayta (720072), Esperanza
(720119), GLKS-58-1642.4 (800290), G-1
(278072.1 O), Mariva (720025), ARK-69.1
(675158), and Gabriela (720120). In San
Ramon, dry matter of the same materials
reached about 22-23%, with the varieties
Dejima (800974), Esperanza (720119), G-1
(278072.1 O), CFJ-69.1 (676002), ARK-69.1
(675158), Yungay (720064), Linea-21,
Linea-34, and Gabriela (720120) being the
highest. The mean dry matter content for
these clones was higher in the arid lowlands of La Malina (20.96%) than in the
hot and humid San Ramon area (18.90%),
confirming the detrimental effects of high
temperatures on this variable previously
reported by Hernandez (1989). However,
sorne genotypes produced higher dry
matter contents under warmer conditions
than cooler ones (Linea 34, Linea 83, and
25.0.--------------~
20.0
~ 15.0
~
e:
10.0
Q)
(!)
5.0
o.o .__T'"'"'"""-T'"'"'"""-T'"'"'"""-,.-......,.-...,,.-...,,-...,.-....,,-...,.-....,.-....,---,.........
13 14 15 16 17 18 19 20 21 22 23 24 25 26
DM(%}
Table 1. Analysis of variance far processing parameters combining two environments; 100 from the
pathogen tested list, group 1.
So urce
Mean squares
d.f
Dry matter
Glucose
Chip color
**
**
**
Environnent (E)
1
14.3111
382.730
42.9549
Repetition (E)
2
0.028
0.00108
0.0001
**
**
**
Genotype (G)
99
12.204
0.71581
2.5054
**
**
**
99
0.51266
GxE
5.6612
0.9978
Error
198
0.2791
0.00019
0.0025
C.V
2.660
0.730
1.830
Mean
19.890
1.905
2.740
Note: ** = P > F < 0.0001.
Mex-750826), and others showed outstanding stability for this character (Yungay, Flor
Blanca, CFE-69.1, Mex-32, and Huaycha).
Analysis of variance indicated a significant level of genotype x environment
(G x E) interaction for dry matter, glucose
content, and chip color (P < 0.0001) (Table
1). This finding was contrary to earlier
reports (Shaw and Booth, 1982).
Glucose content
Frequency distributions for glucose content
(Figure 2) and color of chips (Figure 3) also
showed substantial variability among
clones with a tendency for higher values
in San Ramon, and there were significant
differences between environments. For
example, a higher percentage of genotypes with commercially acceptable
Grades 1 to 3 for chip color occurred in La
Molina (69%) compared to San Ramon
(54%). Severa! varieties showed adequate
50.0
-:
LM
SR
SR
20.0
40.0
gi
~ 30.0
.LM
0
25.0
60.0 . - - - - - - - - - - - - - - - - - - ,
15.0
(/)
Q)
20.0
10.0
Q)
C!l
1.5
2.5
3.5
4.5
5.0
1.5
2.5
3.5
4.5
189
25.0~------------~
16
~
20.0
~
12
Ul
Ul
c.
C!l
(])
e(])
PTL(4x)
Phu-stn (2x)
15.0
(])
-5'
e
(])
C!l
16
17 18
19 20 21 22 23 24 25 26 27 28 29 3l 31
DM (%)
10.0
5.0
15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 3l 31
DM(%)
Materials from CIP's lowland tropics virusresistant population were evaluated over
different seasons at all three test sites. Five
clones were found to be suitable for
processing as french fries, chips, or both.
Table 2 describes a group of successful
progenitors of virus resistance whose
clonal evaluation revealed favorable
postharvest characteristics. The National
Potato Program and Tacna and lea Universities in Peru have released sorne of these
as commercial varieties, including
Y84.027 as Mara Bonita, LT-8 as
Costanera, and C91.612 as Mara Reiche.
Effect of storage conditions
Measurements taken on the harvest from
breeding materials in Huancayo showed
that weight loss during ambient storage
was higher (reaching levels of 8%, the
Table 2. Advanced progenitors with good processing quality and potential for use, La Molina, winter, 1994,
group 3.
Resistan ce
Yield (kg/plant)
DM (%)
GI (%)1
Clone
Use 2
LT-8
PVX, PVY
0.850 (1.35)
20.94 (1.05)
CH,FF
0.05
Y84.027
PW
1.150 (3.84)
18.93 (2.95)
CH,FF
0.05
1.070 (2.30)
20.19 (1.85)
X86.011
PVX
0.06
CH
PLRV, PVX
1.000 (3.09)
88.108
24.38 (1.03)
CH
0.05
PW, PVX
1.200 (2.40)
22.50 (0.92)
C91.612
0.06
FF
Note: DM = Dry matter.
1 GI = glucose content (acceptable below 0.25).
2 CH = chips; FF = french fries.
190
Research on Potato
A
25.0
24.5
24.0
:2
o 23.5
23.0
22.5
30
60
90
Days of storage
B
0.25
0.20
l0.15
3l
8::J 0.10
0.05
0.00
30
60
90
Days of storage
Table 3. Correlations among dry matter content (DM), specific gravity (SG), oil absorption, and glucose
content (GC) of 16 late blight and frost tolerant clones, at harvest and after storage, group 3.
R
Harvest
Ambient storage
Cold storage (4C)
30
60
90
30
60
90
SG-DM
0.912 **
0.725 **
0.880 **
0.928 **
0.613
0.774 **
0.747 **
-0.473
-0.538 *
SG-oil
-0.458
-0.605 *
-0.645
-0.893 ** -0.565
-0.235
-0.622 ** -0.560 *
-0.362
-0.181
-0.181
-0.563
SG-GC
-0.296
-0.420
-0.528 *
-0.303
DM-oil
-0.500 *
-0.804 ** -0.606 *
-0.162
-0.457
-0.124
-0.256
-0.341
-0.537 *
-0.386
DM-GC
Ol-GC
0.350
0.100
0.387
0.026
0.292
0.164
0.727
Note: * = significant (P < 0.05); ** = highly significant (P < 0.01 ).
*
**
*
**
191
Table 4. Yield and chip color of selected clones under two storage conditions, La Malina, winter 1995,
group 3.
Chip color1
Yield
DM (%)
Clone
4C
4C
Ambient
(kg/plant)
Ambient
R
30 days
30 days
90 days
90 days
1.160 a
21.4
1.5
3.5
E86.694
1.0
4.0
2.5
22.8
E86.011
0.909 ab
1.0
1.0
2.0
1.5
1.5
LT-8
0.827 b
20.5
1.5
3.0
1.5
3.5
3.5
21.5
4.0
E86.604
0.777 b
1.0
1.0
4.0
3.0
E86.300
0.765 b
22.2
1.0
3.0
2.0
2.5
2.5
MARI VA
0.745 b
22.4
2.0
3.5
2.0
3.5
3.0
0.743 b
21.7
E86.692
1.0
3.5
1.0
4.5
2.5
22.3
E86.695
0.750 b
1.0
3.5
1.0
3.5
3.0
Duncan test for yield/plant (P < 0.05).
1 Scale used: 1-5, where 1= lightest color (white or pale yellow) and 5= darkest color (dark brown). Colors in the 1-3
range are commercially acceptable. R = reconditioning at ambient temperature following 90 days cold storage.
192
Research on Patato
Table 5. Estimation of combining ability effects using a diallel design with reciprocals, Group 3.
Progenitors
Combining ability effects (GCA)
Yield
SG
RS(1)
RS(2)
378015.16 (TS-2)
168.2
8.2
0.001
-0.006
C83.119
-16.8
-4.2
-0.019
0.023
378017.2 (LT-7)
664.6
3.1
-0.028
-0.032
1-1039
-89.6
-2.5
0.012
0.001
Katahdin
-4.3
-5.0
-0.032
-0.003
377250.7
-70.6
-2.3
0.012
-0.015
377964.5
7.9
0.1
0.014
-0.007
575049 (CEW-69.1)
-59.3
2.6
0.390
0.040
SE (gi)
SE (gi-gj)
25.95
39.94
0.72
1.08
0.016
0.026
0.012
0.018
Note: RS(1 ), measured 10 days after harvest; RS(2), measured 60 days after harvest (ambient).
Discussion
These preliminary evaluations of CIP
populations and advanced clones for
processing quality have identified a useful
range of variabi 1ity and a number of
promising clones with good processing
characteristics. The majority of these also
contain resistance to at least one important disease, such as late blight or
common viruses. Clones with tolerance to
cold storage have also been identified.
193
94
Research on Potato
References
Burton, W.G. 1989. The Patato. 3rd
Edition. Longman Scientific &
Technical, Essex, UK. 742 p.
Dogras, C., A. Siomos, and C. Psomakelis.
1989. Sugar content, dry matter and
sprouting of patato (Solanum tuberosum
L.) tuber stored at 6C and 1OC in
relation to cultivar and area of
production. Patato Abstracts 16(3):114.
FAO. 1995. Potatoes in the 1990s.
Situation and prospects of the world
patato economy. lnternational Patato
Center (CIP), Food and Agriculture
Organization (FAO), Rome, ltaly. 39 p.
Gamble, M.H., P. Rice, and J.D. Selman.
1987. Relationship between oil uptake
and moisture loss during frying of patato
slices from c.v. Record U.K. tubers.
lnternational Journal of Food Science
and Technology (UK) 22:233-241.
Gautney, T.L. and F.L. Haynes. 1983.
Recurrent selection for heat tolerance in
diploid potatoes (Solanum tuberosum
subsp. phureja and stenotr:Jmum).
American Potato Journal 60(7):537-542.
Hernandez, E. 1989. Herencia de los
Factores de calidad para procesamiento
195
The line x tester method is preferred for the evaluation and selection of
parental lines for hybrid true potato seed (TPS) varieties where propagation is
sexual and the hybrid TPS is to be used for raising a commercial potato
(So/anum tuberosum L.) crop. Hence, the line x tester procedure was used to
generate two sets of hybrid TPS families. The first set comprised two standard
male lines from the andigena group (S. tuberosum subsp. andigena) crossed
with 39 new female lines from the tuberosum group (S. tuberosum subsp.
tuberosum) to produce 78 hybrid families. The second set comprised nine new
andigena male lines crossed to six standard tuberosum female lines to produce 54 hybrid families. These hybrid families were evaluated under four
contrasting thermo-photoperiod environments: winter and spring, and summer
and autumn seasons at CIP research stations in La Molina (240 m) and
Huancayo (3280 m), respectively, during 1999-2000. The C1 seedling tubers
were also grown at a late blight (LB) hot spot in Comas (2650 m) to evaluate
their reaction to LB caused by Phytophthora infestans (Mont.) de Bary and the
effect of LB on yield. This is the first study in potato using the line x tester
method and analysis of the data using stability characters for the selection of
parental lines. The results show the approach is useful for selection of potential parental lines for the production of stable hybrid TPS varieties. However,
this selection procedure needs to be validated by multilocation trials in subsequent years with direct seedling transplants from hybrid TPS families
involving the selected female and male lines.
The first evaluation involving a heterozygous tester was suggested in maize
breeding to provide a measure of general
combining ability (GCA) in line x tester
studies, where a group of new lines were
available for testing but there was no plan
for using them in a specific hybrid combination. The lines remaining after selection
for GCA could then be tested further in
single crosses to obtain information on
1
197
198
Research on Potato
Results
The performance of the nine new male
lines for the five characters calculated
from the four trials is given in Table 1. All
the males showed values less than that of
Desire for al 1 five characters. The proportional contributions of the male lines,
female lines, and of the interactions were
also calculated (Table 1). The contribution
of the female testers was much higher for
plant vigor and uniformity, and for marketable yield compared with males and the
male x female interactions. The males
seem to be contributing slightly higher to
the uniformity of tuber shape than the
females, but significantly higher than their
interactions. Si mi larly, the contribution of
the males for total tuber yield was higher
than that of the females but less than the
contribution made by their interactions.
The stability analysis and the simultaneous
selection of the male lines are presented
in Table 2. Only one male, C96LB-13.3,
was selected for al 1 five characters as was
the check Desire, indicating that this
male line performed best with all six
female testers studied. The male line
C94H-07.3 (no. 5) was selected for highest
stability values for plant vigor and unifor-
Males
1
2
3
4
5
6
7
8
g
1o
C95LB-22.7
C96LB-13.3
C96LB-67.4
C96L-12.19
C94H-07.3
C94H-07.6
C95T-07.3
C95Hl-07.3
C95Hl-08.3
Dsire (check)
LSD 0.05
8.1
7.7
7.6
7.3
7.4
8.3
0.1
CV';b
Male (1)
6.6
7.89
Proportional
contribution
Female (t)
(%)
Male x female (1 x t)
1 Scale: 9
Plant vigor1
7.5
7.7
7.5
7.7
14.18
4.21
8.3
2.63
9.61
2.62
8.6
0.24
9.34
5.43
Total y1eld
(Vha)
16.96
18.47
17.42
18.33
18.20
17.03
18.49
19.25
16.07
28.32
0.87
Marketable
yield (%)
70.51
78.28
78.37
77.06
77.67
79.47
73.35
77.15
70.42
96.02
1.57
17.08
6.01
4.13
8.30
7.27
13.27
23.47
7.56
= excellent, 1 = poor.
CIP Program Report 1999 - 2000
199
and thei r i nteractions far the five characters revealed that (1) contributions of the
female lines were the highest far plant
vigor, unifarmity far plant and tuber shape,
and far total yield and (2) the contribution
far marketable yield was highest far the
male testers.
Table 2. Stability comparison far plant vigor, plant uniformity, tuber shape uniformity, total yield, and
marketable yield of new putative males of hybrid TPS families with six standard females evaluated at La
Malina and Huancayo, Peru, 1999 and 2000.
Tuber shape
Total yield
Marketable
Plant vigor
Plant
Male lines
No.
yield (%)
uniformity
uniformity
(Vha)
-1
-3
-1
-5
C95LB-22.7
2
1
C96LB-13.3
2
6+
7+
6+
8+
8+
4
-3
C96LB-67.4
3
4
3
7+
C96L-12.19
2
4
4
9+
5+
5+
-4
C94H-07.3
2
5
11 +
10+
5+
. C94H-07.6
o
o
6
5+
7+
10+
C95T-07.3
4
1
7
5+
6+
7+
C95Hl-07.3
o
o
3
11 +
8
10+
-1
-2
C95Hl-08.3
1
2
3
9
Dsire (check)
10
10+
13+
5+
13+
5+
**
**
**
**
**
Males
**
**
**
**
**
Environments
**
*
ns
ns
ns
1nte ractio n
**
Heterogeneity
ns
ns
ns
ns
**
Residual
ns
ns
ns
ns
0.25025
0.37922
0.28708
Pooled error
9.327
30.597
Notes: + = selected males, * = significant at 0.05, ** = significant at 0.01, ns = non-significant.
Table 3. Evaluation of 9 new male lines of hybrid TPS families with 6 standard female lines (line x tester) as
F1 C1 tuber families far total yield), late blight score (AUDPC), and general combining ability (GCA) at
Comas, Peru, 1999-2000.
No.
Males
Yield (Vha)
GCA
AUDPC
GCA
C95LB-22.7
1
28.57
8.32
736.75
-609.21
C96LB-13.3
2
20.74
0.49
1440.25
94.29
C96LB-67.4
3
13.92
-6.33
1929.08
583.12
C96L-12.19
4
21.88
1.63
1213.33
-132.63
C94H-07.3
5
21.86
1.61
1152.86
-193.10
C94H-07.6
-1.22
19.03
1350.42
6
4.46
7
C95T-07.3
15.29
-4.96
1884.94
538.98
C95Hl-07.3
14.81
-5.44
8
1744.24
398.28
C95Hl-08.3
-5.91
9
26.16
683.86
-662.10
10
Amarilis (check)
39.22
956.67
LSD 0.05
2.38
157.60
se(g)
1.80
119.16
2.55
168.51
se(Q - g.)1
CV%
17.80
17.71
Proportional Males (1)
50.62
60.05
contribution Females (t)
22.69
24.96
Males x Females (1 x t}
15.62
(%}
11.80
AUDPC = area under the disease progress curve.
200
Research on Potato
Table 4. Performance comparison of 39 new female lines of hybrid TPS families with 2 standard males
(line x tester design) evaluated at La Malina and Huancayo, Peru, 1999 and 2000.
Plant vigor1 Uniformity score {avg.)1
Fema les
No.
Total yield
Marketable
yield (%)
{t/ha)
Tuber shape
Plant
C95LB-02.8
7.7
7.6
6.7
18.48
69.25
1
C95LB-03.19
6.7
17.71
2
7.6
7.7
76.23
C95LB-08.3
7.7
7.7
6.7
17.54
81.04
3
C96LB-35.9
6.8
14.03
72.44
7.4
4
7.4
75.62
C96LB-35.B.4
7.3
7.0
6.6
16.32
5
76.62
6.7
16.11
C96LB-35.B.6
7.4
7.4
6
80.41
C96LB-50.B.2
7.5
7.1
19.03
7.6
7
20.74
79.68
C96LB-59.2
6.9
6.8
7.2
8
6.5
15.91
74.70
C96LB-59.12
7.0
6.7
9
6.8
17.74
80.78
C96LB-63.9
7.2
7.3
10
17.80
C96LB-63.B.3
7.0
6.8
6.8
75.50
11
C96LB-68.1
6.8
6.3
15.30
74.08
7.2
12
16.93
80.16
6.8
C96L-34.4
8.0
7.8
13
79.07
6.7
17.90
C96L-34.7
8.0
7.9
14
6.8
15.96
77.18
C96L-35.3
7.3
7.5
15
7.2
18.37
74.26
C96L-45.6
7.6
7.3
16
7.1
20.33
77.24
C96L-46.3
7.7
7.4
17
16.61
C96L-57.1
7.8
7.5
6.7
77.40
18
C97L-01.3
6.8
7.0
16.78
77.87
7.2
19
7.0
6.9
17.56
79.41
C97L-01.4
7.1
20
6.8
18.17
72.93
C97L-04.2
7.4
7.2
21
19.08
79.06
7.7
7.2
C96T-02.1
7.6
22
76.34
6.9
19.62
C96T-02.9
7.9
7.8
23
6.7
16.89
78.73
C95H2-02.2
7.8
7.6
24
18.59
76.62
6.8
7.6
C95H3-16.4
7.7
25
6.9
18.77
76.86
7.8
C95H3-16.5
7.8
26
6.8
17.07
76.24
C95H3-19.1
7.7
7.5
27
6.8
16.92
76.14
C95H3-19.11
7.4
7.5
28
6.8
16.68
73.29
C95H3-19.13
7.7
7.6
29
7.0
17.16
75.49
C95H3-19.16
7.5
7.6
30
19.12
77.57
7.9
6.9
C95HL-08.1
7.8
31
6.8
15.67
73.66
7.6
C95HA-01.2
7.4
32
7.2
18.17
71.94
7.5
7.5
C96H-01.3
33
17.41
76.90
7.6
6.9
C96H-01.6
7.6
34
17.81
73.32
6.9
C96H-02.4
7.4
7.6
35
21.04
76.33
7.0
C96H-02.7
8.0
7.7
36
7.1
16.61
77.13
7.7
C96H-04.7
7.8
37
6.9
17.46
71.76
7.6
C96H-08.8
7.7
38
6.6
16.32
73.77
7.6
C96H-10.2
7.4
39
27.16
96.72
8.5
7.9
Dsire tuber (check)
8.1
40
0.3
1.42
2.58
0.3
LSD 0.05
0.2
8.2
16.52
6.91
cv (%)
6.7
8.5
Proportional contribution (%)
13.66
6.40
12.79
10.6
Female (1)
11.09
0.72
27.33
0.15
0.14
0.16
Male (t)
5.49
1.27
2.30
1.36
1.74
Female x male {I x t}
1 Scale: 9
= excellent, 1 = poor.
201
Discussion
The development of parental lines poses
no problem equal in complexity to that
involved in the evaluation of lines. The
final evaluation of even the most carefully
selected lines must rest upon their performance in hybrid combinations. At CIP,
Mendoza (1985) studied GCA of 20 highyielding clones mated to two testers using
the line x tester model and evaluated
them under three environments for yield,
tuber uniformity, and seedling survival. He
suggested that to use TPS in commercial
potato production, it is important to
identify parental clones with high GCA for
those three characters. However, wel 1defined approaches for the evaluation and
selection of parental clones were lacking.
The studies reported by Upadhya and
Cabello (1997) marked the beginning of
202
Research on Patato
Table 5. Stability comparison for plant vigor, plant uniformity, tuber shape uniformity, total yield, and
marketable yield of new females of hybrid TPS families with two standard males evaluated at La Malina
and Huancayo, Peru, 1999-2000.
Tuber shape
Total yield
Plant
Marketable
Plant vigor
No. Female lines
uniformity
uniformity
yield (%)
(t/ha)
1 C95LB-02.8
-2
3
26+
29+
32+
2 C95LB-03.19
15
9
22+
30+
23+
C95LB-08.3.
3
5
28+
31 +
41+
19+
-2
15
12
2
10
4 C96LB-35.9
7
4
3
6
14
5 C96LB-35.B.4
4
12
11
5
6 C96LB-35.B.6
20+
18
9
36+
27+
39+
7 C96LB-50.B.2
16
4
3
8 C96LB-59.2
36+
36+
-1
-10
o
-5
11
9 C96LB-59.12
6
9
16
10 C96LB-63.9
20+
40+
o
o
12
11
C96LB-63.B.3
15
20+
-1
o
3
1
9
12 C96LB-68.1
14
16
12 C96L-34.4
38+
38+
36+
8
20+
34+
14 C96L-34.7
40+
40+
2
17
9
21+
27+
15 C96L-35.3
6
4
38+
31+
16 C96L-45.6
21+
8
17 C96L-46.3
26+
37+
39+
28+
11
5
19
30+
18 C96L-57.1
33+
9
5
2
33+
31+
19 C97L-01.3
17
1
5
24+
C97L-01.4
35+
20
3
14
7
11
29+
21
C97L-04.2
36+
33+
22+
32+
38+
22 C96T-02.1
19
C96T-.02.9
28+
38+
23
37+
34+
4
9
20
32+
24
C95H2-02.2
38+
33+
20+
24+
20+
25 C95H3-16.4
30+
34+
25+
27+
34+
36+
26 C95H3-16.5
14
16
10
16
27 95H3-19.1
32+
13
16
12
12
18
C95H3-19.11
28
10
5
16
27+
29 C95H3-19.13
30+
15
8
16
23+
32+
C95H3-19.16
30
18
25+
31+
C95HL-08.1
34+
40+
31
7
1
11
24+
C95HA-01.2
32
2.0+
1
22+
17
38+
C96H-01.3
20+
33
17
24+
24+
C96H-01.6
22+
34
22+
4
25+
10
28+
C96H-02.4
24+
35
16
34+
41+
40+
32+
36 C96H-02.7
4
26+
35+
C96H-04.7
34+
37
36+
-4
18
C96H-08.8
27+
28+
38
29+
8
2
7
8
24+
C96H-10.2
39
35+
39+
43+
43+
Dsire (check)
34+
40
**
**
**
**
**
Females
**
**
**
**
**
Environments
*
**
*
ns
ns
lnteraction
*
**
ns
ns
ns
Heterogeneity
*
**
ns
ns
ns
Residual
27.842
8.435
0.31851
0.4014
0.5072
Pooled error
Notes: + = selected females, * = significant at 0.05, ** = significant at 0.01, ns = non-significant.
203
Table 6. Evaluation of new female lines of hybrid TPS families with 2 standard male lines (line x tester) as F1
C1 tuber families far total yield, late blight score (AUDPC), and general combining ability (GCA) at Comas,
Peru, 1999-2000.
GCA
AUDPC
GCA
Female Lines (no.)
Yield (t/ha}
No.
825
-181
C95LB-02.8
20.80
2.24
1
-343
22.11
5.55
663
C95LB-03.19
2
-2.69
1041
35
C95LB-08.3
15.87
3
-7.96
1215
209
10.60
4
C96LB-35.9
-5.45
1443
437
C96LB-35. B.4
13.11
5
-5.92
1314
12.64
308
C96LB-35.B.6
6
-224
24.84
6.28
782
7
C96LB-50.B.2
879
-127
C96LB-59.2
26.40
7.84
8
-0.12
-151
C96LB-59.12
18.44
855
9
13.64
-4.92
1456
450
C96LB-63.9
10
-2.24
1470
464
C96LB-63. B.3
16.32
11
-5.10
13.46
1561
555
12
C96LB-68.1
20.28
1.72
868
-138
13
C96L-34.4
20.91
2.35
926
-80
C96L-34.7
14
-5.08
C96L-35.3
13.48
1196
190
15
-4.95
C96L-45.6
13.61
1176
170
16
-4.21
C96L-46.3
14.35
1230
224
17
C96L-57.1
15.01
-3.55
988
-18
18
-4.68
C97L-01.3
13.88
1594
588
19
-5.36
C97L-01.4
13.20
1352
346
20
-4.96
1444
C97L-04.2
13.60
438
21
C96T-02.1
24.41
5.85
668
-338
22
-126
C96T-.02.9
20.83
2.27
880
23
C95H2-02.2
23.94
4.62
755
-251
24
20.29
1.73
957
-49
C95H3-16.4
25
21.14
2.58
770
-236
C95H3-16.5
26
416
-590
24.09
5.53
27
C95H3-19.1
-0.08
1173
167
C95H3-19.11
18.48
28
10.28
-671
C95H3-19.13
28.84
335
29
18.03
-0.53
959
-45
C95H3-19.16
30
15.38
-3.18
1183
177
C95HL-08.1
31
691
-315
C95HA-01.2
21.89
3.33
32
26.68
8.12
609
-397
C96H-01.3
33
-326
21.86
3.25
680
34
C96H-01.6
C96H-02.4
26.43
7.87
537
-469
35
28.55
9.99
554
-452
C96H-02.7
36
-7.73
C96H-04.7
10.83
1282
276
37
11.29
-7.27
C96H-08.8
1507
501
38
-3.53
C96H-10.2
15.03
1019
13
39
Amarilis (check)
34.46
1126
40
3.94
271
LSD 0.05
1.99
137
se@)
2.82
se@- Q)1
194
18.6
CV%
24
AUDPC = area under the disease progress curve.
204
Research on Patato
Conclusions
The results of this study show that the line
x tester approach is useful for selecting
parental lines from a group of new lines
available for testing, but without any plan
for their use in a specific hybrid combination design for the production of hybrid
TPS varieties. However, this selection
procedure has to be validated by conducting multilocation/year trials with direct
seedling transplants from hybrid TPS
families involving the selected female and
male lines.
References
Bradshaw, J.E. and G.R. Mackay. 1994.
Breeding strategies for clonally
propagated potatoes. In: Bradshaw,
J.E. and G.R. Mackay (eds.). Potato
genetics. CAB lnternational,
Wallingford, UK. p. 467-498.
Kang, M.S. and R. Magari. 1995. STABLE:
A basic program for calculating stability
and yield-stabi 1ity statistics. Agronomy
Journal 87:276-277.
Mendoza, H.A. 1985. Selection of uniform
progenies to use TPS in commercial
205
206
Research on Patato
207
208
Research on Potato
for the characters studied were nonsignificant (Table 1 ), hence the data were
analyzed individually for seed size and
density classes. The results of the analysis
for the effect of seed density and seed size
on the characters are presented in Tables
2a and 2b. Seed density seems to have
sorne effect on emergence but does not
affect plant vigor and total and marketable
yields (Table 2a). Seed size, however,
significantly affected emergence and total
and marketable yields (Table 2b). Plant
vigor does not seem to be affected by seed
size or density at 45 days after transplanting. There is a highly significant correlation
between seed size and emergence
(r = 0.78, P <0.01) and between seed size
and marketable yield (r = 0,44, P <0.01 ),
but only a significant correlation between
seed size and total yield (r = 0.50, P <0.05).
No significant correlation was found
between seed size and plant vigor (r = 0.38,
P >0.05), or between seed size and plant
uniformity
(r = 0.22, P >0.05).
Simmonds (1963) reported higher germination percentages (85-90%) for large and
medium seeds compared with small seeds
(60-65%). Dayal et al. (1984) showed that
the 1000 TPS weight in a hybrid TPS
family was positively correlated with total
yield. Later, Thakur and Upadhya (1994)
reported a highly significant effect of seed
size on seedling emergence and growth in
nursery beds. Seedlings grown from large
seeds had higher survival and total tuber
yield. The higher yield was due to the
production of more tubers and higher
yield/plant. They also showed that seed
size had a positive but non-significant
relationship with average tuber weight and
marketable yield. In the present study,
however, there was a significant effect of
seed size on marketable yield. This could
be due to the fact that Thakur and
Upadhya (1994) studied the effect of seed
size in the open pollinated TPS of TPS-2,
whereas the present study used the hybrid
TPS family from Serrana x TS-5. This
difference could be due to the effect of
hybrid vigor.
Table 1. ANOVA far seed density and size on seedling emergency, plant vigor, total yield, and marketable
yield as seedling transplant crop of Serrana x TS-5 hybrid TPS family at la Malina, 1999.
ANOVA
Emergence DAS 9 days
Plant vigor
Total yield (Vha)
Marketable yield (%)
MSD 0.05
Density(D)
ns
ns
ns
*
Size(S)
ns
**
**
*
DxS
ns
ns
ns
ns
C\1%
2.9
9.0
9.50
2.2
Notes: * = significant at 0.05; ** = significant at 0.01; ns = not significant.
Table 2a. Comparison of seed density on seedling emergence, plant vigour, total and marketable yield in
Serrana x TS-5 hybrid TPS family as seedling transplant crop at la Malina, 1999.
Density
Emergence at 9 days
Vigor 45 DAT
Total yield Vha
Marketable tubers
(%)
(%)
7.6a
17.61a
01
91.5a
80a
88.2b
16.77a
7.5a
02
79a
C\1%
2.9
9.0
9.50
2.2
Notes: Duncan at P= 0.01. 01 =High density; DAT =Days atter transplant; D2=Low density; Vigor: 1=Poor, 9=Excellent.
Table 2b. Comparison of seed size on seedling emergence, plant vigour, total and marketable yield in
Serrana x TS-5 hybrid TPS family as seedling transplant crop at la Malina, 1999.
Seed size
Emergence at 9 days
Vigor 45 DAT
Total yield Vha
Marketable tubers
(%)
(%)
19.2a
8.0a
81.0a
94.4a
S1
79.3b
16.8b
91 .8a
7.4a
S2
16.0b
78.6b
7.2a
83.4b
S3
9.0
9.50
2.2
2.9
C\1%
Notes: Duncan at P = 0.01.
S1: > 1.6 mm; S2: 1.6-1.4 mm; S3: 1.4-1.27 mm.
DAT =Days after transplant; Vigor: 1=Poor, 9=Excellent.
Conclusions
The present study suggests a strong correlation between seed size and yield.
CIP Program Report 1999 - 2000
209
References
Bhatt, A.K., T.C. Bhalla, H.O. Agrawal,
M.O. Upadhya, and N. Sharma. 1988.
Effect of seed size on imbibition and
germination of open pollinated true
seeds of patato. Seed Research 16:1 78182.
Bhatt, A.K., T.C. Bhalla, H.0. Agrawal,
and M.O. Upadhya. 1989. Effect of seed
size on protein and lipid contents,
germination and imbibition in true
patato seeds. Patato Research
32:477-481.
Oayal, T.R., M.O. Upadhya, and
S.N. Chaturvedi. 1984. Correlation
studies on 100 true seed weight, tuber
yield and other morphological traits in
patato (Solanum tuberosum L.). Patato
Research 27:185-188.
Kang, M.S. and R. Magari. 1995. STABLE:
A bas~c program for calculating stability
and y1eld-stability statistics. Agronomy
Journal 87:276-277.
Khatana, V.S., M.O. Upadhya, A. Chilver,
and C.C. Crissman. 1996. Economic
impact of true patato seed on patato
production in Eastern and North-Eastern
India. In: Walker, T.S. and C.C.
Chrissman (eds.). Case studies of the
economic impact of CIP-related
technologies. lnternational Patato
Center, Lima, Peru. p. 139-156.
Lowe, LB. and S.K. Res. 1972. Effects of
environment on the relation between
seed protein and seedling vigor in
wheat. Canadian Journal of Plant
Science 52:1 57-164.
Lowe, LB., G.S. Ayers, and S.K. Ries.
1972. Relationship of seed protein and
amino acid composition to seedling
vigor and yield of wheat. Agronomy
Journal 64:608-61 O.
Pallais, N. and R. Falcan. 1997.
Temperature and moisture affect
dormancy and deterioration of true
21 O
Research on Patato
Between 1993 and 1999, new hybrid progenies of true potato seed (TPS) were
adopted by 100,000 small-scale farmers on 3500 ha in Vietnam, or about 10%
of total potato (Solanum tuberosum L.) area. Most adoption occurred in the
Red River Delta in northern Vietnam during the winter crop season. Hybrid
TPS is estimated to have increased potato yield by an average of 6.8 t/ha, or
75%, compared with old varieties grown from clonal seed tubers. Aggregate
economic benefits from TPS in Vietnam are estimated at about US$1 million
per year. The net present value of the investment in TPS research and extension over 1990-201 O is estimated to be between $0.25 million and $2.92
million, yielding a rate of return to research of 29-42%. TPS is estimated to
have increased net household income of adopters by $11.00/year, or 1.2%.
New sources of improved clonal seed tubers, such as imported seed from
China and new improved Vietnamese varieties, may limit further diffusion of
TPS in Vietnam.
2 Vietnam
and other plant pathogens can be transmitted through tubers to the next generation,
subsequently reducing plant health and
yield. Using TPS avoids both of these
limitations, because no portian of the
useable harvest needs to be diverted for
seed and diseases are much less prevalent
in the botanical seed compared with
vegetatively propagated material (Sadik,
1983). These factors have generated
considerable interest in TPS, especially for
use by poor, limited-resource farmers in
developing countries.
Many of the field applications of TPS so
far, however, have not lived up to this
promise. lt has been difficult to achieve
quality and efficiency in both the production and handling of TPS. Consequently,
the cost of TPS to farmers is not insubstantial and the seed itself is sometimes of
uncertain quality. Furthermore, few
farmers have been able to obtain an
economically viable crop from TPS itself.
GIP Program Report 1999 - 2000
211
212
Research on Potato
Table 1. Patato production costs, yields, and economic returns by source of seed in the Red River Delta,
Vietnam.
Seed source
TPS nurseries
TPS
Old local clonal lmported clones
New local
and seedling
seedling
variety
from China
clonal variety
transplants1
tubers
(Ackersegen)
(Mira mostly)
(KT-3)
Input quantity (per ha)
Labor (h/ha)
5243
4569
4094
4094
3814
684
1162
Seed (kg/ha)
0.119
1162
1396
TPS seedlings/ha
72716
14.5
Manure (t/ha)
17.7
13.4
13.4
16.3
N (kg/ha)
120
133
136
136
179
84
125
125
P (kg/ha)
86
83
72
82
42
K (kg/ha)
77
82
Farm-supplied inputs (US$/ha)
Land
50
50
50
50
50
Labor
749
653
585
545
585
43
Man ure
61
33
33
39
Total farm-supplied
860
745
667
667
633
inputs
Purchased inputs (US$/ha)
o
o
o
Nursery plastic cover
8
o
102
328
256
256
Seed
308
101
116
116
Total chemical fertilizer
103
105
11
14
1
16
Total pesticide
1
444
428
Total purchased inputs
224
373
373
Output
11.25
8.99
12.50
19.90
Average yield (t/ha)
15.79
6:30:50:15
36:30:27:7
45:39:16:0
47:27:20:6
39:39:20:2
A:B:C:D (%) 2
Average price of
95
85
95
95
104
yield (US$/t)
Economic returns (US$/ha)
1191
2077
Gross value of yield
1070
1336
857
Net income 3
846
893
484
818
1649
1016
Economicprofit4
(13)
147
(184)
151
Source: Farm surveys in Red River Delta during 1999/00 winter season.
TPS = True patato seed.
1 An average of 0.119 kg of TPS sown in 215 m2 nursery beds is required to produce 72, 716 one-month-old TPS
seedlings. The seedlings are then transplanted to 1 ha.
2 Percent size distribution of yield. A = large (> 100 g/tuber), B = medium (50-100 g/tuber), C = small (20-50
g/tuber), O = very small (< 20 g/tuber).
3 Net income is defined as the gross value of production minus the cost of purchased inputs.
4 Economic profit is defined as the gross value of production minus the cost of all purchased and farm-supplied inputs
(land, labor, and manure).
213
214
Research on Potato
research and extension. The other scenarios result in higher estimates of net
present value because there continue to be
benefits from TPS between 2001-201 O.
Under scenario 3, where TPS benefits are
assumed to continue at a constant level
between 2001 and 201 O, the net present
value at the 10% discount rate is US$2.97
million. The rate of return to research and
extension is 28.6% under scenario 1 and
around 40% under scenarios 2 and 3.
215
Table 2. Benefit-cost analysis of true patato seed (TPS) research and extension in Vietnam (constant
US$'000}.
Aggregate net benefits
lnvestment in
Net economic benefits
Year (season)
to farmers 1
research and
to Vietnam 1
extension
Scen. 2
Scen. 3
Scen. 1
Scen. 2
Scen.3
Scen. 1
(78)
o
o
o
78
(78)
(78)
1990/91
o
78
(78)
(78)
o
o
(78)
1991/92
o
o
(92)
(92)
o
92
(92)
1992/93
o
o
(105)
(105)
o
105
(105)
1993/94
(126)
(126)
132
(126)
5
5
1994/95
5
28
185
(157)
(157)
(157)
28
28
1995/96
13
198
198
198
185
13
13
1996/97
1052
1237
1237
1237
185
1052
1052
1997/98
185
790
976
976
976
790
790
1998/99
1075
1075
1075
185
889
889
889
1999/00
o
967
1075
185
(185)
782
889
2000/01
o
147
928
860
1075
(147)
713
2001/02
o
752
1075
147
928
(147)
606
2002/03
o
645
1075
134
(134)
510
940
2003/04
o
537
1075
134
(134)
403
940
2004/05
o
430
1075
93
(93)
337
982
2005/06
o
322
1075
93
982
2006/07
(93)
230
o
215
1075
2007/08
93
(93)
122
982
o
107
1075
93
2008/09
(93)
15
982
o
o
2009/10
1075
93
982
(93)
(93)
o
o
o
78
2010/11
(78)
(78)
(78)
Present value
1526
2869
10% discount rate
4073
1100
426
1769
2972
15% discount rate
1037
1786
2370
787
250
1584
999
Benefit-cost ratio
1.39
3.70
2.61
(10% discount rate)
Benefit-cost ratio
1.32
2.27
3.01
(15% discount rate)
Interna! rate of return,
28.6%
38.9%
41.6%
1990-201 o
1 Scenario 1: no TPS benefits after 2000; Scenario 2: TPS benefits decline to Obetween 2001 and 201 O; Scenario 3:
216
Research on Potato
Acknowledgments
The authors would like to thank Tom
Walker for his suggestions and Elske Van
de Fliert for kindly providing needs
assessment farm survey data for this study.
References
Chilver, A., T.S. Walker, V.S. Khatana, H.
Fano, R. Suherman, and A. Rizk. 1999.
On-farm profitabi 1ity of true potato seed
(TPS) utilization technologies. Social
Science Department Working Paper No.
1999-3. lnternational Potato Center,
Lima, Peru. 41 p.
Fuglie, K., Do Thi Bich Nga, Dao Huy
Chien, and Nguyen Thi Hoa (2001 ). The
economic impact of True Potato Seed in
Vietnam. In: Fuglie, K. (ed.). Performance and Prospects of Hybrid True
Potato Seed in South and Southeast
Asia. Proceedings of the CIP-ADB
Symposium, Field-Testing Hybrid TPS in
the Lowland Tropics of Asia, held at
Bogor, Indonesia, September 13-14,
2000. CIP-ESEAP, Bogor, Indonesia.
p. 151-176.
Sadik, S. 1983. Potato production from true
seed-present and future. In: Hooker,
W.J. (ed.). Research for the potato in the
year 2000. lnternational Potato Center,
Lima, Peru, p. 18-25.
217
Since its founding in 1971, CIP has invested considerable resources in potato
(So/anum tuberosum L.) genetic improvement. CIP's contribution to varieties
released by national programs and to varieties grown by farmers was assessed from survey data in 30 countries that account for 85 % of
developing-country production. In the 1990s, about 40% of national releases
were directly related to CIP's activities. However, by 1997, CIP-related
releases only accounted for about 6 % of potato-growing area. CI P-related
materials have had wider acceptance in smaller national programs with less
area planted to potatoes than in stronger national agricultura! research
system (NARS). Moreover, CIP-related varieties are still relatively young
because varietal change is substantially slower in potatoes than in cereals
and in most other major field crops. In spite of this seemingly disappointing
performance in adoption, we calculate that the rate of return on investing in
potato genetic improvement at CIP is positive at 15-17%.
Shortly after the founding of CIP, a planning conference was held to establish
priorities and recommend specific programs for the use of the wealth of genetic
resources becoming available to the
institute (CIP, 1974). A few high priority
recommendations, such as improving the
content and quality of protein, were
subsequently dropped and sorne low
priority recommendations, i.e., the use of
sexual propagation by means of true
potato seed, were later emphasized. Sorne
research areas, such as adaptation to very
warm growing regions and resistance to
bacteria! wi lt, caused by Ralstonia
solanacearum, have proven to be technically difficult. Others, such as resistance
to late blight (LB), caused by Phytophthora
1
219
Varietal Release
Data on varietal release are usually the
most available information on the performance of a plant breeding program.
However, release does not imply adoption
nor is it a necessary condition for success.
The release of Kerr's Pink in Kenya in 1927
marked the first variety listed in the 30country sample. Between 1927 and 1998
about 500 varieties were released. The first
CIP-related variety was released in 1979.
The pace of varietal release has accelerated since the 1950s, when on average
two varieties were released per year across
the 30 countries. During the 1980s and
1990s, average release peaked at 1 7/year.
This rising and then stabilizing trend of
varietal releases reflects the pattern of
investment in public-sector NARS (Alston
et al., 1 998). In many publ ic-sector NARS,
growth in the early 1960s to the early
1980s was followed by a period of stagnation and even decline in public-sector
spending for agricultura! research.
The institutional source of the germplasm
on which released varieties are based can
be divided into three categories: 1)
developing country NARS, 2) CIP, and 3)
220
Research on Potato
Varietal Adoption
The data on varietal adoption show a very
credible performance by NARS-bred
material (Table 3). About 50% of the
released varieties belong to this category,
and they accou nted for 60% of area
planted to potatoes in the 30-country
sample in 1997.
Shares of released varieties and area
coverage were also congruent for developed country clones with about one
released variety in four and one hectare
planted in four. This correspondence attests
From 1990-98
38.0
32.4
1.1
1.7
2.8
17.6
41.5
7.4
30.7
3.4
2.8
2.8
176
221
attributed to farmers.
Research on Potato
Table 4. _Back of the envelope calculation on returns to CIP's investment in patato breeding.
Assumpt1ons
Period
50 years, 1972-2021
Area
1997, no trend assumed
Prices
Constant (1992) (US$11 O/t)
Source of benefit
Yield increase of 2.5 Vha = US$220/ha
CIP costs
Real; 55% of expenditure on patato
Seed, extension, and ali other research costs
US$11 O/ha
Net benefit
US$110/ha
Results
Adoption ceiling in 2021 (%)
IRR1
NPV 2 (millions US$)
5.8
15
39
10.0
16
51
15.0
17
71
1
IRR
Conclusions
Proportionally and even absolutely, CIP's
impact has been greatest in small NARS.
The adoption of CIP-related patato material in developing countries has also been
modest compared with the performance in
wheat, rice, and maize. Nonetheless, we
show a solid rate of return on investment
of about 15% in CIP's patato breeding
activities. The lack of a very attractive
rate of return to breeding does not imply a
mediocre performance for a patato crop
improvement program as a whole. For a
high-value, vegetatively propagated crop,
there is potential to add value through
223
Acknowledgments
About 60 NARS and CIP scientists contributed to this project. We are particularly
grateful to Jim Bryan, Charles Crissman,
Peter Ewell, Fernando Ezeta, Francisco
Flores, Osear Hidalgo, Hilario Da Silva
Miranda Filho, and Yi Wang. We thank the
lmpact Assessment and Evaluation Group
of the Consultative Group on lnternational
224
Research on Patato
References
Alston, J.M., P.G. Pardey, and
J. Roseboom. 1998. Financing agricultura! research: lnternational investment
patterns and policy perspectives. World
Dev. 26(6):1057-1071.
CIP (lnternational Patato Center). 1974.
Strategy for Utilization. Report of the
lnternational Potato Center's Planning
Conference on Uti 1ization of Genetic
Resources. Lima, Peru. 85 p.
CIP (lnternational Potato Center). 1998.
Working Document on CIP Plant
Breeding Strategies for Patato and
Sweetpotato. Interna! ly Commissioned
Externa! Review (ICER). Lima, Peru. 27 p.
FAO (Food and Agriculture Organization
of the United Nations). FAOSTAT
1998(November). [http://apps.fao.org].
Walker, T.S. 1994. Patterns and implications of varietal change in potatoes.
Social Science Department Working
Paper, No.1994-3. CIP, Lima, Peru. 40 p.
225
226
Research on Potato
227
Research on Patato
Results
lnteractions of genotype, environment,
and management
The performance of three varieties with
different levels of resistance (Amarilis,
Yungay, and Tomasa) was assessed under
three levels of fungicide protection over
3 yr. Yield decreased significantly with
increasing LB disease (regression slope =
-14.9; df = 1, 299; F = 32.8; P < 0.001 ).
Relative AUDPC decreased significantly
with an increase in the frequency of
fungicide treatment (regression slope =
-0.92; df = 1, 356; F = 14.8; P < 0.001 ).
There was an increase in yield with an
increase in the frequency of fungicide
treatment (regression slope = 28.1;
df = 1, 335; F = 11.27; P < 0.001), but
there was a much stronger effect of variety
as indicated by the F value (regression
slope = 77.8; df = 1, 335; F = 110.8; P <
0.001 ). Furthermore, a more complex
model, where both fungicide and variety
were treated as qual itative variables and
including two- and three-way interactions,
also revealed that variety (df = 2, 125;
F = 277.5; P < 0.001) hada much stronger
effect on yield than fungicide treatment
(df = 2, 125; F = 76.5; P < 0.001 ).
Two of the two-way interactions were
significant for yield: fungicide treatment x
environment (df = 46, 125; F = 6.35;
P = 0.001) and GxE (df = 46, 125;
F = 8.46; P < 0.001 ). In contrast, the
interaction genotype x fungicide treatment
was not as significant (df = 4, 125;
F = 2.50; P = 0.046). The results of
ANOVA for rAUDPC were similar to those
for yield. The main effects of variety,
fungicide, and environment on rAUDPC
were all highly significant, as were all
two- and three-way interactions (df =
2-94, 131; F ~ 2.01; P < 0.001). When
individual rAUDPC values were regressed
against the mean rAUDPC of the three
genotype observations at the correspondi ng site-year, the effect of GxE was strong,
but no cross-over effects were present
(Figure 1 A). In contrast, the interaction
0.8
0.6
0.4
a"'
0.2
C1
11
CI
O l-~~.ti~!::::li~t:::[:::!:=-:_~_:!-~!_!_~_H~__!E~~ci~~e!Bj
1
B
Amarillis
0.8
0.6
Clci
CI
0.4
CI
CI
..... o
!I
0.2
a..
..
:::>
~
~
e:
0.8
Yungay
0.6
0.4
0.2
0.8
0.6
0.4
0.2
0.1
0.2
0.3
0.4
0.5
0.6
Figure 1. lnteractions of variety x environment (A) and culivar x environment x fungicide (B-D). Mean relative
area underthe disease progress curve (mean rAUDPC) for all genotypes (Amarilis, Yungay, and Tomasa) was
used as an index of disease pressure.
CIP Program Report 1999 - 2000
229
A set of 16 released varieties and advanced breeding lines was tested in the
FPR-FFSs. Since the variety x environment
interaction was significant, along with the
main effects for both yield and rAUDPC
(df ~ 16, 786; F ~ 4.95; P < 0.001 ), joint
linear regression was performed (Table 1).
Chata Roja showed the highest overall
yield. The mean yield of Chata Roja
(16.4 kg/plot (ca. 27 t/ha); n = 48) was
9.1 times higher than the mean yield of
the most susceptible entry, Canchn
(1.8 kg/plot (ca. 3 t/ha); n = 35). Chata
Roja, Amarilis, and Kory showed relatively
high yields irrespective of the environmental index for yield (mean yield for three
selected genotypes) or disease pressure
(mean rAU DPC for three selected genotypes). Susceptible entries, including two
of the advanced breeding lines tested,
showed low yields at all environments,
and yields were negatively correlated with
disease pressure. Among the varieties with
mean yields over 1O kg/plot, i.e., the more
resistant entries, only Atahualpa showed
yields that were significantly negatively
correlated with rAUDPC.
The most promising entry identified
through the 1997/98 FPR-FFS was Amarilis,
a LB-resistant variety with which few
farmers in the area were previously
familiar. After its success in the FPR-FFS,
CARE provided credit to allow larger-scale
production of Amarilis, and it has rapidly
gained acceptance in the area. For
example, 35% of families who participated in FPR-FFSs had commercial
plots with Amarilis after two cropping
seasons of participation, compared with
10% of nonparticipating families. This
10% suggests that if a FFS group finds a
good variety or breeding line, it will
230
Research on Potato
Table 1. Stability parameters far released varieties and advanced breeding line entries over 1997-1999. Regression coefficient (b, slope) plus its standard error (SE)
are given far entry yield regressed on disease-pressure index (mean rAUDPC far Amarilis, Yungay, and Tomasa at the corresponding site) and entry yield regressed
on ~eld environmental index {mean ~ield far Amarilis, Yunga~. and Tomasa at the corres~onding site}. lndex ranges are shown to indicate com~arabili!Y of results.
Entry
Yield (kg/plot)
RAUDPC 2
Yield regressed on mean
Yield regressed on mean
lndex ranges
yield
rAUDPC
3 p4
p4
Meanmean
b SE
Mean yield
Mean rAUDPC
n
b SE
n
0.074
0.77 0.24
0.002
9.27 6.20
0.144
3.8-16.3
0.07-0.53
16.4 a
Chata Roja
48
36
0.107
0.59 0.19*
0.002
3.96 4.61
1.0-16.3
0.07-0.59
14.3 ab
0.392
Amarilis
84
90
0.056
0.61 0.20*
0.002
7.02 4.97
0.07-0.59
Kory
14.2 ab
0.162
1.0-16.3
84
90
0.168
1.14
0.18
<0.001
-0.56
5.49
0.920
1.0-16.3
0.07-0.59
Chagllina
13.1
be
81
72
0.042
1.19
0.22
10.20
5.71
3.8-16.3
13.1
be
<0.001
0.083
0.07-0.53
Cipira
48
36
0.132
1.35
0.27
<0.001
-3.66
8.02
0.651
3.8-16.3
0.07-0.53
12.9 be
Victoria
48
36
0.086
10.25
6.93
1.0-11.1
0.11-0.59
1.26 0.32
<0.001
0.146
12.4 be
66
377740.2
54
0.186
-22.07
15.29
1.71 0.30*
<0.001
0.165
4.7-16.3
0.07-0.40
Mara Tambea
12.0 be
21
27
-12.73 3.78
1.0-16.3
0.160
1.16 0.10
<0.001
0.001
0.07-0.59
11.9 be
90
Atahualpa
84
0.227
1.04 0.14
<0.001
-7.77 4.32
0.077
1.0-16.3
0.07-0.59
10.1 e
78
Perricholi
72
0.002
-5.154.56
1.0-5.8
0.11-0.59
0.169
1.25 0.38
0.267
7.7 d
54
Jerusaln
36
0.56 0.13*
-7.19 3.54
1.0-16.3
0.07-0.59
0.335
<0.001
0.046
Yungay
5.8 d
90
84
0.26
0.002
-10.81 3.04
1.0-8.7
0.11-0.59
0.62 0.19
0.001
4.6 ef
57
Libertea
47
0.375
1.07 0.25
-11.30 2.69
1.0-5.8
<0.001
<0.001
0.11-0.59
3.8 ef
52
380076.1
36
0.103
7.9-7.9
0.07-0.07
3.0 ef
Molinera
3
3
0.567
0.36 0.07*
<0.001
-9.66 1.61
1.0-16.3
0.07-0.59
2.2 f
<0.001
To masa
83
83
-6.17 3.03
1.0-16.3
0.451
0.95 0.25
<0.001
0.050
0.11-0.59
1.8 f
60
Canchn
35
C")
""C
1 CIP
=e
a
'
~
;>
"C
o
;::::.
"'
"'
"'1
N
o
o
o
""~
(l.)
N
;;;
CI>
"'
CI>
~
::r
o::::1
'"O
o~
Table 2. Stability parameters far true patato seed (TPS) trials and new breeding lines tested in three FFS during the 1999/2000 season. Regression eoeffieient (b,
slope) plus its standard error (SE) are given far entry yield regressed on disease-pressure index (mean rAUDPC far Amarilis, Yungay, and Tomasa at the
eorresponding site) and entry yield regressed on yield environmental index (mean yield fr Amarilis, Yungay, and Tomasa at the eorresponding site). lndex ranges
are shown to indieate eomEarabili~ of results.
Entry1
lndex ranges
Yield regressed on mean
Yield regressed on mean
Yield (kg/plot)
rAUDPC 2
yield
rAUDPC
p~
p~
Mean3
b SE
Mean }'.ield Mean rAUDPC
mean
b SE
n
n
True Patato Seed
0.07-0.40
0.546
10.05 13.52
0.470
0.9-10.2
7.7 a
0.055
0.27 0.43
16
18
6
0.0-10.2
0.07-0.63
0.73 0.24
0.006
-1.34 6.76
0.845
24
6.4 ab
26
0.074
7
0.0-10.2
0.07-0.63
0.147
0.75 0.22
0.002
-6.94 6.08
0.268
22
5.9 abe
24
1
0.0-10.2
0.07-0.63
1.40 0.21
-14.77 8.24
0.088
Yungay
22
5.8 abe
23
0.291
<0.001
0.0-10.2
0.07-0.63
1.12 0.18
-9.88 6.80
0.161
24
5.7 abe
26
0.155
<0.001
Amarilis
0.295
0.0-10.2
0.13-0.63
5.6 be
20
0.135
1.23 0.21
<0.001
-8.29 7.66
18
5
-0.82 5.41
0.881
0.0-10.2
0.07-0.63
24
0.110
0.61 0.18*
0.002
22
5.0 be
4
0.07-0.63
0.56 0.15*
0.001
-3.65 4.53
0.429
0.0-10.2
24
3.9 ed
26
0.134
2
0.474
0.0-10.2
0.07-0.63
26
0.124
0.43 0.13*
0.004
-2.75 3.77
24
3.9 ed
3
0.0-10.2
0.48 0.11 * <0.001
-6.04 3.37
0.087
0.07-0.63
24
2.3 d
26
0.383
Tomasa
Group 1 New breeding lines
0.029
1.09 0.53
0.111
-18.9 90.7
0.846
14.8-27.6
0.04-0.15
6
27.0 a
6
Cipira
14.8-27.6
0.04-0.15
26.1 a
0.016
0.65 0.30
0.095
-25.9 51.1
0.639
391683.29 (11)
6
6
0.200
14.8-27.6
0.04-0.15
24.3 ab
0.093
0.86 0.25
0.026
-72.0 46.9
393079.24 (32)
6
6
-140 68.1
0.108
14.8-27.6
0.04-0.15
22.5 ab
6
0.023
1.47 0.36
0.016
391011.17 (19)
6
0.129
14.8-27.6
0.04-0.15
0.78 0.40
0.123
-92.5 48.5
19.3 b
6
0.106
393248.55 (40)
6
0.120
-157 57.4
0.052
14.8-27.6
0.04-0.15
13.4 e
0.274
1.150.58
Perrieholi
6
6
Group 2 New breeding lines
-75.3 47.4
0.163
4.1-23.4
0.02-0.18
0.062
1.32 0.23
0.001
21.0 a
8
391137.7 (10)
8
0.022
4.1-23.4
0.02-0.18
0.074
1.26 0.22
0.001
-103 33.7
17.7 ab
8
393077.54 (31)
8
0.71 0.37
0.100
-15.8 44.4
0.734
4.1-23.4
0.02-0.18
15.4 be
8
0.046
Cipira (2)
8
4.1-23.4
0.02-0.18
-55.9 29.0
0.103
0.067
0.88 0.13
<0.001
13.8 be
8
393371.159 (47)
8
4.1-23.4
0.02-0.18
0.008
-126 35.5
0.012
12.6 e
8
0.116
1.35 0.35
392639.31 (26)
8
0.02-0.18
-48.5 15.6
0.021
4.1-23.4
0.078
0.61 0.08
<0.001
12.5 e
8
393339.242 (44)
8
0.004
-72.7 20.5
0.012
4.1-23.4
0.02-0.18
11.7 e
0.139
0.81 0.18
8
393385.39 (51)
8
0.003
-110 11.5 <0.001
4.1-23.4
0.02-0.18
8
0.213
1.05 0.22
10.8 e
Perrieholi
8
*b signtticantly different from b = 1.0 (Eberhart and Russell, 1966) in yield response.
1 CIP breeding program genotype number or Peruvian variety name with FFS synonym in parentheses. lf bolded, the entry was in the top 4 chosen from 26 by farmer groups in San Miguel.
2 rAUDPC is the ratio of AUDPC to the total area of the graph (Fry, 1978).
3 Means followed by same letters are not signtticantly different at 0.05 leve! according to SNK multiple-range test.
4 Probability that
slope equals O.
Discussion
The FPR-FFSs have provided farmers the
opportunity to evaluate and gain access to
varieties previously little known in the
area, as well as to breeding lines not yet
formally released. The farmer groups were
enthusiastic about several of the varieties,
including older promising lines and newly
identified breeding lines. Amarilis, which
was tested in FPR-FFSs in San Miguel
starting in 1997, has become increasingly
popular in the San Miguel area. Chata
Roja, which was tested starting in 1998,
was the most outstanding entry that year
and was subsequently released by
UNHEVAL. Two additional breeding lines
were released by other institutions after
successful testing in the FPR-FFSs, while
two others were rejected dueto poor
performance. A set of new breeding lines
and a set of TPS-derived populations were
tested through the FFSs, and further testing
is now in progress.
The primary purpose of the genotype x
management experiments was to demonstrate that fungicide spray decisions should
be predicated both on the level of resistance of the crop, and on the environment.
233
E':S
.:,..
en
m
a
::r
o::::
"C
g.
Table 3. Stability parameters far a group of new breeding Hnes tested in tour FFSs during the 1999/2000 season. Regression eoeffieient (b, slope) plus its standard
error (SE) are given far entry yield regressed on disease-pressure index (mean rAUDPC far all entries tested at the eorresponding site) and entry yield regressed on
yield environmental index (mean yield far all entries tested at the eorresponding site). lndex ranges are shown to indieate eomparability of results.
rAUDPC2
Entry1
Yield {kg/plot)
Yield regressed on mean
Yield regressed on mean
lndex ranges
yield
rAUDPC
!!a.
o
391580.30 (21)
Cipira
387212.41 (9)
393371.157 (46)
393073.197 (29)
393280.57 (41)
392633.54 (24)
Perrieholi
380011.12 (1)
n
8
8
8
8
8
8
8
8
8
Mean 3
23.3 a
21.5 ab
21.0 abe
20.8 abe
19.4 abed
18.4 abcd
17.1 bcd
15.8 ed
14.8 d
n
8
8
8
8
8
8
8
8
8
mean
0.053
0.010
0.009
0.039
0.026
0.075
0.039
0.196
0.089
b SE
1.07 0.16
1.05 0.17
1.12 0.13
1.24 0.19
0.99 0.16
1.010.17
0.87 0.21
0.92 0.23
0.73 0.17
p4
<0.001
<0.001
<0.001
<0.001
<0.001
<0.001
0.006
0.007
0.005
b SE
3.87 205
-148 193
-108 204
-135 232
4.63 191
-107 190
-244 152
-264 159
-215 122
p4
0.986
0.471
0.616
0.580
0.982
0.594
0.161
0.147
0.128
mean yield
6.1-25.3
6.1-25.3
6.1-25.3
6.1-25.3
6.1-25.3
6.1-25.3
6.1-25.3
6.1-25.3
6.1-25.3
mean rAUDPC
0.03-0.07
0.03-0.07
0.03-0.07
0.03-0.Q?
0.03-0.07
0.03-0.07
0.03-0.07
0.03-0.07
0.03-0.07
*b significantly different from b = 1.0 (Eberhart and Russell, 1966) in yield response.
1 IP breeding program genotype number or Peruvian variety name with FFS synonym in parentheses. lf bolded, the entry was in the top 4 chosen from 26 by farmer groups in San Miguel.
2 rAUDPC
is the ratio of AUDPC to the total area of the graph (Fry, 1978).
Means followed by same letters are not significantly different at 0.05 leve! according to SNK multiple-range test.
4 Probability that slope equals O.
235
Acknowledgments
We gratefully acknowledge the IFAD
and the OPEC Fund far lnternational
Development far their financia! support
far fieldwork. Data analysis was supported
by a U.S. Agency far lnternational Development linkage grant from CIP to
K.A. Garrett, J.J. Heath, and J.L. Heath.
We would like to thank F. Ezeta and
A. Gonzales (CIP), and F. Boeren (CARE)
far their important contributions in the
early planning stages of this work. We
thank G. Thiele and R. Hijmans (CIP) far
helpful discussions and far critica! readings
of the manuscript.
References Cited
Ceccarelli, S., S. Granda, R. Tutwiler,
J. Baha, A.M. Martini, H. Salahieh,
236
Research on Potato
68:1650-1655.
Landeo, J.A., M. Gastelo, H. Pineda, and
F. Flores. 1995. Breeding far horizontal
resistance to late blight in patato free of
R genes. In: Dowley, L.J., E. Bannon,
L.R. Cooke, T. Keane and E. O'Sullivan,
(eds.). Phytophthora infestans 150.
European Association far Patato
Research by Boole Press, Ltd., Dublin,
lreland. p. 286-274.
Matteson, P.C. 1996. lmplementing IPM:
Policy and institutional revolution.
Journal Agricultura! Entomology
13:173-183.
Nelson, R., R. Orrego, O. Ortiz, J. Tenorio,
C. Mundt, M. Fredrix, and N.V. Vien.
2001. Working with resource-poor
farmers to manage plant diseases. Plant
Disease 85(7):684-695.
Ortiz, O., P. Winters, and H. Fano. 1999.
La percepcin de los agricultores sobre
el problema del tizn tardo o rancha
(Phytophthora infestans) y su manejo:
Estudios de casos en Cajamarca, Per.
Revista Latinoamericana de la Papa
11 :97-120.
SAS lnstitute. 2000. SAS Online Doc, SAS/
STAT User's Guide, release 8.1 Edition.
SAS lnstitute lnc., Cary, NC, USA.
Sperling, L., M. Loevinsohn, and
B. Ntabomvura. 1993. Rethinking the
farmer's role in plant breeding: Local
bean experts and on-station selection in
Rwanda. Experimental Agriculture
29:509-519.
Thiele, G. 2000. Bibliography of
participatory research in CIP. Social
Science Department Working Paper No.
237
238
Research on Patato
2 CIP-UPWARD,
3
239
240
Resean:h on Potato
Table 1. Comparison of original rice Farmer Field School (FFS) and the emerging patato integrated disease
management (IDM) approach in Nepal.
Aspect
Rice FFS
Patato IDM
Remarks
Multi-season
Time frame
Season-long
IDM requires a longer time trame since its
success is determined by doing a followup by replanting produced seeds in next
seasons.
Learning plots Experimentation
Experimentation, seed Seed is an important component of IDM.
Learning plot is also used to multiply/
multiplication/
maintain good-quality seed.
maintenance
Weekly, but with more Depends on appearance of disease
Frequency of Weekly
symptoms, especially far late blight.
frequent inspection
sessions
Sessions need not be weekly early in the
far late blight
season, however, they need to be
detection.
more frequent (2-3 per week) when
late blight/ bacteria! wilt symptoms begin
to appear.
To be used more selectively since weekly
AESA needs to be
Basic method far
AESA 1
AESA produces data which may not be
complemented by
learning by
directly usefuVrelevant far patato IDM.
other 'discovery'
'discovery' by
methods
farmers
Directly and indirectly Unlike insects, pathogens are often not
Making things Directly through
visible. Experiments to show the 'effects'
visible
AESA
needs to be done.
Disease management takes several seasons
1mpact after severa!
lmpact after FFS
Evaluation
to complete. lmpact assessment needs to
seasons
season
be done only after several seasons.
Disease and seed management are closely
Multiple constraints Single constraint Scope
interrelated. FFS needs to deal with the
cropping system
crop
interaction among disease and seed
factors, as well as dynamics between
patato and other crops.
1 AESA =
agro-ecosystem analysis.
241
Crop management
X
Table 3. Profile of FFS participants and their change in knowledge on patato IDM.
Change in knowledge
Farmers gender
FFS Sites
Literacy (%)
Pre-test
Post-test score Net increase
(M/F)
(%)
score (%)
(%)
31
70
18 (18/0)
83
39
Sarlahi
57
27
84
Rupandelhi
34 (31/3)
40
78
40
34(21/13)
82
38
Sunsari
Nigaley
73
38
24 (8/16)
50
35
Danushi
25 (17/8)
60
38
98
60
Surkhet
39 (3/36)
60
73
18
Kathmandu
21 (18/3)
100
55
32
53
85
Kavre
25 (21/4)
90
20
65
85
Kapilbastu
17 (14/3)
100
43.8
80.8
37
Total average 237/26 (17/9)
74
1
242
Research on Potato
Difference
(%)1
79.5
211.1
105.3
108.6
157.9
32.7
60.4
30.8
84.5
Yield increase
(Vha)
27.5
14.0
15.0
5.8
22.0.
8.0
10.0
2.5
14.2
Difference
(%)2
220.0
116.7
75.0
72.5
61.1
47.1
40.0
9.2
80.2
Percent of difference compares yield in FSP plots and increase in yield obtained in IDM plots.
243
References
Bhomi, B.K. 1997. Potato development
programme in Nepal: Past, present
and future. In: Pradhanang, P.M. and
J. G. Elphingstone (eds.). lntegrated
management of bacteria! wi lt of potato:
Lessons from the hills of Nepal. Lumle
Agricultura! Research Centre, Pokhara,
Nepal. p. 4-10.
Hidalgo, O. 1998. Major diseases and the
control components available far their
utilization in IDM programs in countries
of SWA. Paper presented at the
Workshop on the lmplementation of
lntegrated Disease Management
Program Utilizing the Farmer Field
School Approach held 2-5 June in
Kathmandu, Nepal. (unpublished.)
Lama, T.L. 1998. Potato development
programme in Nepal. Paper presented at
the Workshop on the lmplementation of
lntegrated Disease Management
Program Utilizing the Farmer Field
School Approach held 2-5 June in
Kathmandu, Nepal. (unpublished.)
MNASD (Ministry of Agriculture/
Agricultura! Statistics Division). 1999.
Statistical information on Nepalese
244
Research on Potato
Described here are the evolution through the years of the tuber-seed production efforts of the Nepal Potato Program in cooperation with CIP and efforts
to promote the institutional establishment of the Seed Producer Groups (SPG)
in the country.
Nepal is cine of poorest countries in the
world, with an annual growth rate of 2.4%.
lts population reached 23 million in 2000.
About 90% of the people live in rural
areas and are mainly dependent on
agriculture. Their food needs are becoming
critica! as the population grows.
Potato plays an important role in meeting
diese food needs. Per capita consumption
of potato in Nepal is 30 kg/year (CIP,
1998), slightly higher than the world
average of 28 kg/year. In 1999, total area
under potato was 118,043 ha and productivity was 9.2 t/ha (MNASD, 1999).
However, yield is much lower than that in
neighboring countries and the world
average (16 t/ha). The major cause for this
low productivity is poor quality tuber-seed,
non-availability of good quality seed, and
lack of knowledge about modern crop
cultivation methods (Ojha, 1999).
Nepalese farmers have cu ltivated potato
for 200 years and potato is one of the
major food crops in the mid- and high-hills
where it is a staple food. This differs from
the plains and urban areas where potato is
mainly considered a vegetable. Potato
could play a more important economic
role if it could be produced in a sustainable manner. This begins with good quality
tuber seed.
1
2
3
245
246
Research on Potato
----.
i
Group formation
process
Group management
----.
i
Seed potato
production
Seed marketing
lnternal quality
control mechanism
_.
Figure 1. Operational steps and achievements for the organization and implementation of the SPG in Nepal.
247
Table 1. Comparison of tuber-seed produced by Seed Producer Groups with farmers seed in three
Nepal. Winter and summer 1998, and winter 1999.
Region
Mid-Western
Far-West
Central
Yield Samples
Yield Samples
Yield Samples
Vha
(no.)
Vha
(no.)
t/ha
(no.)
Terai, winter 1998
24.1
27.41
19.97
131
85
90
1 Seed groups (Basic seed)
14.8
21.94
25 1
20
15.20
17
2 User groups (lnspected seed)
-3
0.0 3
14.40
15
9.20
35
3 Non-groups (Farmer seed)
90.3
117.10
Yield lncrease (%) (1 vs. 3)
Hills, summer 1998
27.90
147
12.25
30 2
90
23.56
1 Seed groups (Basic seed)
-2
20.55
O.O
30
22.70
30
2 Users groups (lnspected seed)
7.95
3 Non-groups (Farmer seed)
9.21
35 2
30
10.90
36
33.0
180.40
Yield lncrease (%) (1 vs. 3)
116.20
Terai, winter 1999
1 Seed groups (Basic seed)
33.2
45
20.0
20.0
90
55
2 Non-groups (Farmer seed)
14.3
45
7.9
45
40
13.5
132.2
153.4
48.5
Yield lncrease {%) {1 vs. 2)
regions of
Average
yield
(Vha)
23.25
17.79
10.76
116.10
23.75
21.63
9.38
153.20
23.1
12.1
90.9
248
Research on Patato
-e
~-~
The SPG approach is a sustainable farmerrun program. This approach has shown
greater opportunity to expand the seed
production industry in Nepal because this
approach is simple, cost effective, resultoriented, and viable in small-farmer
communities. Based on this experience the
Ministry of Agriculture has approved SPG
as a tool for technology dissemination to
produce good quality seed in the country.
lt is expected that the SPG approach will
continue to be implemented by farmers in
a self-sustainable manner. This approach
could be implemented for informal quality
seed production of other crops with certain
modifications.
We recommend that the socio-economic
impact of the SPG approach be assessed.
O,.....
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References
CIP (lnternational Potato Center). 1998.
Potato Facts: A compendium of key
figures and analysis for 32 important
potato-producing countries. Lima, Peru.
249
250
Research on Potato
Project CIP/SDC. 1998. Annual report Informal high quality seed potato
production and marketing by Seed
Producer Group in Nepal. Potato
Development Project for Bhutan, Nepal
and Pakistan (Project CIP/SDC-Nepal)/
Potato Research Programme, Khumaltar,
La 1itpu r, Nepal.
Project CIP/SDC. 1999. Annual report
informal high quality seed potato
production and marketing by Seed
Producer Group in Nepal. Potato
Development Project for Bhutan, Nepal
and Pakistan (Project CIP/SDC-Nepal)/
Potato Research Programme, Khumaltar,
Lalitpur, Nepal.
Thapa, B., D. N. Ojha, B. Saha, and O.A.
Hidalgo. 1999. Manual to guide Seed
Producer Groups for the implementation
of rules and regulations for seed patato
quality control. Potato Research
Program/Project CIP/SDC, Khumaltar,
Lalitpur, Nepal. 17 p.
2
3
for seed directly, but subsequent generations of certified seed used and distributed
by farmers increase its impact on the
informal seed system and on patato
production. Strengthening links between
them would improve the whole patato
seed system (Thiele, 1999; Aguirre et al.,
1999).
PROINPA ran a series of trials in three
potato-producing regions of Bolivia from
1990 to 1996 to evaluate the performance
and profitability of certified seed in
farmers' fields over several multiplications. Results of these trials were gathered
and then analyzed in 1999 and are
reported here. The objectives of this
research were to (1) study the agronomic
performance of certified seed compared to
farmers' seed, (2) assess the relative
profitabi 1ity of certified seed over the
study period, and (3) evaluate the conditions that affect economic performance.
251
Evaluation factors
The trials were carried out in three potatogrowing areas in the highlands above 3000
m where recurrent frost and drought limit
potato production (Table 1). Certified seed
was compared to farmers' seed under their
own crop management practices. The
trials were implemented with small
farmers (< 1 ha of potato). Soil fertility was
generally poor with low organic matter
content (1 .0%) and phosphorus (< 1 O ppm),
and adequate potassium (> 0.4 meq/100 g).
Farmers usually applied 5-7 t/ha of sheep
or cattle manure to their potato fields and
low levels of chemical fertilizers, usually
in the form of NPK 18-46-0 or urea according to cash availability.
Seed Origins
Two native commercial varieties
(S. tuberosum andigena) were used,
Waycha and Sani lmilla. The certified
seed was obtained from a semiprivate
company UPS/SEPA, the main provider of
high quality seed to the formal seed
system. The farmers' seed had different
origins; these are discussed in the results
section. The first year the same quantity of
seed (24 kg) for each seed source was
planted with each farmer in two randomized plots with two replications. In
following years, the subsequent generations of each seed source were planted
following the same design using the same
quantity of seed for each seed source.
The two different seed sources were
compared for several years, four years in
Cochabamba, six years in Chuquisaca,
and five years in Tarija.
Table 1. Altitude, average climatic conditions, and variety used in the regions where the trials were carried
out, Bolivia, 1990-96.
Region/Department
Altitude
Annual temperature
Annual rainfall
Variety
(m)
(C)
(mm)
used
Cochabamba1
3490
9
531
Waych'a
Chuquisaca2
3105
12
537
Sani lmilla
Tarija2
3300
12
325
Sani !milla
1 Source: Meteorological Station form Experimental Research Station, Toralapa.
2 Source: National Meteorological Service (SENAMHI).
252
Research on Potato
Results
Farmers' seed source
1990
1991
1992
1993
1994
1995
1996
Years
253
254
Research on Potato
Economic analysis
Two sources of economic benefits related
to yield were identified: (1) yield differences, and (2) price differences. The price
differences are attributed to variation in
the distribution of tuber size between
farmers' own seed and certified seed
managed under their conditions. Higher
yields are strongly associated with more
remunerative prices because of the greater
proportion of higher priced, larger-sized
tubers.
The initial cost of certified seed of Bs.
1.3/kg (US$1.00 = Bs.3.17) was much
higher than farmers' seed of Bs.0.40/kg,
which was estimated from the ware patato
market price. For the later multiplications,
the same price was al located to the
progenies of both seed types.
In the three regions, for farmers with
complete records, certified seed gave both
higher yields and was sold at higher
average prices than farmers' saved tuber
seed. On average, the yield difference
5CXXl
400J
400
:DXl
2CXXJ
350
1000
300
o
250
f fJ
t t
-1000
200
-2000
150
-3000
-4000
100
-5000
Tarija
Chuquisaca
Cochabamba
1
1234
year
Cochabamba
1234
1234
year
Chuquisaca
year
Tarija
255
256
Research on Potato
-2000
10
20
30
40
Yield (tlha)
Conclusions
These data support the notion that certified
seed was moderately profitable in two of
the three regions where it was the subject
of repeated on-farm testing in the 1990s.
The increases in yield due to certified seed
vary from farmer to farmer and are relatively modest on average because of
constraints such as viruses, insects, and
adverse climatic conditions, especially in
Chuquisaca and Tarija. In Cochabamba, it
seems the general farmers' crop management practices, including seed quality,
were better in the area where the trials
were carried out.
From the data it would appear that the
variation in the difference in gross benefits
Acknowledgements
The authors would like to thank Enrique
Fernandez Northcote and Vctor Alvarez
for their efforts in the virological tests and
analyses for these trials.
References
Aguirre, G., J. Calderon, D. Buitrago,
V. lriarte, J. Ramos, J. Blajos, G. Thiele,
and A. Devaux. 1999. Rustic seedbeds,
a potential bridge between formal and
traditional potato seed systems in
Bolivia. In: lmpact on a changing
world: Program report 1997-98. CIP,
Lima, Peru. p. 195-204.
Terrazas, F., V. Suarez, G. Gardner,
G. Thiele, A. Devaux, and T. Walker.
1998. Diagnosing potato productivity in
farmers' fields in Bolivia. Social
Science Department Working Paper No.
1998-5. lnternational Patato Center,
Lima, Peru. 71 p.
Thiele, G. 1999. Informal patato seed
systems in the Andes: Why are they
important and what should we do with
them? World Development 27(1 ):83-99.
257
The majority of people in Bangladesh live below the poverty line on an income of less than US$1 per day. lncreased productivity of food crops, such as
potato (Solanum tuberosum L.), would help to alleviate poverty. Farmers can
increase potato productivity by using good quality seed, but poor potato
farmers can afford to huy only poor quality seed from informal sources. Seed
produced through the formal certified seed system meets only 5-6% of the
seed requirement. Potato growers and extension workers are not familiar
with improved agro-techniques for on-farm production of high-quality seed.
Strengthening technology for on-farm seed production and diffusion through
the informal seed system could increase income and enhance the social status
of poor farmers.
259
Methods
Rangpur District in northern Bangladesh
(Figure 1) was identified as the area with
the agro-climatic conditions most suitable
for potato seed production. The area under
potato in Rangpur is about 15% of the total
patato area of Bangladesh. A rapid rural
appraisal was u sed to gather i nformation
from farmers on farmers' seed sources,
number of generations of seed used from
different sources, seed quality, varieties
used by farmers, disease incidence, yields
260
Research on Potato
Results
More than 80% or more of the farmers in
Rangpur grow patato as a cash crop during
winter (November-March). Comparatively
rich farmers buy certified seed from the
BADC and use this seed and a part of their
subsequent production for three or four
generations. Most resource-poor farmers
buy degenerated seed from cold store
owners, farmers, and traders. Cardinal, a
pink-skinned variety, is the main HYV
grown by farmers in Rangpur and other
northern districts. A few farmers plant
Diamant, Kufri Sindhuri, and Heera
varieties. Diamant is more popular in the
central and southern districts, like
Munshiganj near Dakha. Other varieties
such as Multa (obtained from Munshiganj
District) were seen on sorne farms, but the
produce was diseased with common scab
and bacteria! wilt.
Traditional varieties, introduced about a
century ago, were the only potatoes grown
in Bangladesh until 1960 when HYVs were
introduced (Siddique, 1991 ). They are still
widely grown and, although they are lowyieldi ng, bring premium prices above
those of Cardinal and Diamant. They are
still popular among growers and consumers, primarily because of their good
keeping quality in farm potato stores,
relatively low costs of production, reasonably high yields with low inputs and under
stress, and the high market demand due to
better taste. The area grown to traditional
varieties is, however, decreasing as more
.,.
BANGLADESH
kIO O
10 ZO
40
lnternational Bo11ndary
District Boundory
Capital
1 NDIA
( WEST
IENGAL)
BAY
OF
BENGAL
1.1
111
.,.
261
Research on Potato
Price
(US$/kg)
28-40
40-55
28-40
40-55
0.22
0.21
0.38
0.33
28-40
40-55
0.17
0.16
28-40
40-55
0.31
0.27
Discussion
lncreases in production of breeder seed by
TCRC and certified seed by BADC to meet
the requirements of a large number of
resource poor farmers is not possible under
present circumstances of limited resources
and inadequate infrastructure. This is partly
because the lack of extension personnel
leaves a serious void in public sector
support to future seed production programs.
Also, present linkages between the private
sector, non-government organizations, the
BADC, and research and extension
organizations are almost nonexistent.
The formal seed production system of
multiplication is lengthy and takes 6 or 7
yr to reach farmers. Therefore, improving
Conclusions
Research strategies to strengthen the
formal and informal seed production
systems include the following.
Develop strong relationships between
the publ ic sector, private sector, and
strong NGOs for the production and
diffusion of healthy seed of improved
potato varieties.
Provide intensive training on improved
agro-techniques at seed production sites
for seed/ware potato production to
farmers, extension personnel, NGOs,
and the private sector. Farmer field
schools for integrated pest management
presently operated by developmental
organizations and NGOs in Bangladesh
would be useful for disseminating proper
seed flow management in the informal
seed system.
Reduce seed multiplication generations
from six to two at government seed
farms, followed by two multiplications
in farmers fields. After two multiplications at the farm level, farmers can sel 1
the produce as certified seed to other
farmers.
263
References
BBS (Bangladesh Bureau of Statistics).
1998. Statistical pocketbook of
Bangladesh, Bangladesh Bureau of
Statistics, Statistics Division, Ministry of
Planning, Dhaka, Bangladesh. 432 p.
BBS. 1999. Statistical pocketbook of
Bangladesh, Bangladesh Bureau of
Statistics, Statistics Division, Ministry of
Planning, Dhaka, Bangladesh. 466 p.
Choudhury, A.R. 1990. Role of cold stores
in the improvement of seed patato. In:
Proceedings of the lnternational
Seminar on the Seed Patato held from
8-1 O January in Dhaka, Bangladesh.
p. 133-138.
Hossain, M., B.M. Lal, and A. Chowdhury.
2001. Changes in agriculture and
264
Research on Potato
Research on Sweetpotato
Scientist and Farmer
Partners in Research far the 21 st Century
267
Research on Sweetpotato
'
reverse: TCCACAACATATTATTCCCC)
allowed the amplificati o n of a 277-bp
segment corre spondin g to th e coding
seq uence of the oryzacystatin gene.
Eightee n of th e 25 tran sge ni c lines showed
no signs of infection after being inocul ated
with SPFMV-RC-infected grafts from
l. setosa sc ions, whereas the nontran sfo rm ed Jonathan control developed
symptoms of vein clearing and chlorotic
spots, which are typi ca l of SPFMV-RC
infect ion (Figure 1 ). Thi s suggests a
putative res istan ce in the 18 tran sge ni c
1in es.
Seventeen of the asymptomatic transgen ic
lines gave negat ive res ults with NCMELISA, which co nfirmed the visual
observations and demonstrated a high
degree of ag reeme nt between the v isu al
269
setosa stock.
setosa stock.
symptoms in /. setosa after grafting healthy /. setosa scions onto the SPFMV-RC-inoculated transgenic lines.
in /. setosa after grafting healthy /. setosa scions onto the SPFMV-RC-inoculated transgenic lines.
4 NCM-ELISA
270
Research on Sweetpotato
Acknowledgements
The authors wish to thank Dominique
Michaud of Laval University, Canada, for
providing the OCI gene construct and
technical advice. This work was partially
supported by CGIAR-Canada Linkage Fund
(CCLF) Project #05-02-RT.
References
Bevan, M.W., S.E. Masan, and P. Goelet.
1985. Expression of tobacco mosaic
virus coat protein by a cauliflower
mosaic virus promoter in plants
transformed by Agrobacterium. EMBO
Journal 4:1921-1926.
Blandenvoorde, M.F.J., H.S. Brand,
Y.M.C. Henskens, E.C.I. Veerman, and
A.V.N. Amerongen. 2000. Protease
inhibitors in health and disease
control-Medica! and industrial aspects.
In: Michaud, D. (ed.). Recombinant
protease inhibitors in plants. Landes
Bioscience, Georgetown, TX, USA.
p. 202-213.
Carey, E.E., R.W. Gibson, S. Fuentes,
M. Machmud, R.O.M. Mwanga,
G. Turyamureeba, L. Zhang, D. Ma,
F. Abo El-Abbas, R. El-Bedewy, and
L.F. Salazar. 1999. The causes and
control of virus diseases of sweetpotato
in developing countries: Is sweetpotato
virus disease the main problem? CIP
Technical Progress Report (1997-1998).
p. 421-428.
Cipriani, G., D. Michaud, F. Brunelle,
A. Golmirzaie, and O.P. Zhang. 1999.
Expression of soybean proteinase
inhibitor in sweetpotato. lmpact on a
changing world. Program Report 19971998. lnternational Patato Center, Lima,
Peru. p. 271-277.
Clark, C.A. and J.W. Moyer. 1988.
Compendium of sweetpotato diseases.
The American Phytopathological
Society, Minnesota, USA.
Daines, R.H. and W.J. Martin. 1964.
Russet crack, a new virus disease of
271
273
274
Research on Sweetpotato
co2
Analysis of data
All data sets were analyzed by standard
ANOVA procedures for a randomized
complete block design, using the PROC
UNIVARIATE procedure of SAS software
(SAS, 1990).
Table 1. Shoot dry matter, fibrous root dry matter, and storage root dry matter accumulation of two
sweetpotato genotypes at 126 DAT grown in pots with six N levels. Values followed by common letters do
not differ significantly (capital letters refer to N levels; lowercase letters to genotypes).
N1
O.O
0.4
0.8
1.2
1.6
2.0
Mean*
Shoot dry matter (g/plant)
Jewel
5.97 aD
11.04 bC 13.81 bB
14.73 bB
25.93 bA
27.69 bA
16.53
Tanzania 10.31 a F 21.21 a E 39.88 a D 47.26 a e
52.94 a B
71.84 a A
40.58
Mean** 8.14 F 16.12 E 26.85 D 30.99 e
39.44 B
49.78 A
CV[%]
14.80
R2
0.97
6.81 b AB
Jewel
5.50 a BC
7.27 bA
4.52 be
5.61 b BC
6.28 b AB
6.00
Tanzania 9.19 a E 14.67
16.25 a e 13.91 a D
21.58 a A
17.88 aB
15.58
9.76 e
14.20 A
Mean** 7.35 D 10.97 BC 10.39 e
12.08 B
CV[%]
24.05
R2
0.86
Jewel
12.99 be
56.09 bA 58.98 b A 56.51 b A
31.87 b B
36.94 a B
42.23
Tanzania 14.22 a E 67.08 aB 63.72 a B 75.20 a A
34.90 aC
22.02 b D
46.19
Mean** 13.60
cv [%]
61.59
R2
61.35
65.86
33.38
29.48
b
a
b
a
b
a
25.07
0.82
1 Application of N (g/plant).
CIP Prograrn Report 1999- 2000
275
0.25
~Ol
~11/
0.2
'5,
.e
c.
im
~ :::::1/
--
,k.
...
.A
/: .. -~-
~ 0.15
eo
2
1.8
1.6
1.4
,.A
A:
0.1
Jr ""
:e
1.2 "'E
1
o
---
Chorophyll Jewel
-liD- Chorophytl Tanzania
_...... SLN Jewel
SLN Tanzania
0.05
-A-
0.4
1.2
0.8
1.6
0.8
0.6
0.4
0.2
N supply g/plant
00
a
"'O
fJJ
O Tanzania
rl
00
"'O
Jewel
70
b
b
() 40
ri""
rf
b
3)
2)
10
B
., B
0.8
0.4
1.2
1
1.6
N supply g/plant
276
Research on Sweetpotato
Table 2. Discrimination against 13C0 2 [%o] at 126 DAT in leaves of two sweetpotato genotypes grown in
pots with six N levels. Values followed by common letters do not differ significantly (capital letters refer to
N levels; lowercase letters to genotypes).
e lsotope in leaves
N1
o.o
0.8
1.6
0.4
1.2
2
Mean*
-31.11 b e -32.07 b D -31.47 be -30.83 b B -30.12 b A -30.03 b A -30.97 b
Jewel
-30.95 a E -30.88 a E -30.40 a D -29.52 a e -29.01 a B -28.71 a A -29.84 a
Tanzania
-31.26 D -31.44 D -30.83 c -30.09 B -29.52 A -29.30 A
Mean**
-0.36
cv [%]
R2
0.99
Note: a = Mean separation within rows for genotypes by MSD
columns for N levels by MSD = 0.26; P = 0.05.
1 fertilization (g N/plant).
Jewel
O Tanzania
"!/!.
E 15
3'
a
10
O)
g:
:2:
0.4
0.8
1.2
1.6
N supply g/plant
12
OJewel
Tanzania
rf
o 0.8
rl-
~ 0.6
E
~
$:
0.4
b a
f
b
bt
02
. e
0.4
.,
'
0.8
1.2
'
1.6
'
N supply g/plant
277
Conclusion
Results of the pot experiment showed that
both storage root initiation and storage root
growth are delayed with high N supply,
confirming observations of earlier investigators (Lowe and Wilson, 1974;
Villagarcia, 1996). High NAR of Tanzania
confirms the assumption that varietal
differences in photosynthetic efficiency
may depend on both leaf inclination and
chlorophyll content in leaves. A low
extinction coefficient allows more efficient 1ight transfer through canopies. More
vertically inclined leaves might be
advantageous, particularly under high light
conditions by minimizing the probability
of photoinhibition and increasing light
penetration to lower leaves, thereby
maximizing whole-canopy photosynthesis
(Lambers et al., 1998). Consistently higher
NAR values of Tanzania as compared with
Jewel could be attributed to more efficient
light interception of this genotype due to
its more erect leaves. Tanzania also had
higher chlorophyll content per unit leaf
area, which is characteristic for crop
species or varieties adapted to high
irradiance (Lambers et al., 1998).
ldeotypes such as Tanzania may therefore
be more efficient in total plant biomass
production under conditions of high
irradiance, which was the case in San
Ramn.
Concerning the N effect on WUE, it has
been shown that N stress leads to increased stomatal opening, thereby
reducing WUE. Varietal differences in
278
Research on Sweetpotato
References
Farquhar, G.D. and R.A. Richards. 1984.
lsotopic composition of plant carbon
correlates with water-use efficiency of
wheat genotypes. Austalian Journal of
Plant Physiology 11 :539-552.
Hahn, S.K. and Y. Hozyo. 1984. Sweet
potato. In: Goldsworthy, P.R. and
N.M. Fisher (eds.). The physiology of
tropical field crops. John Wiley & Sons,
Ltd, New York, USA.
Hill, W.A., H. Dodo, S. K. Hahn,
K. Mulongoy, and S. O. Adeyeye. 1990.
279
Effect of GxE lnteraction on Root Yield and Betacarotene Content of Selected Sweetpotato (lpomoea
batatas (L) Lam.) Varieties and Breeding Clones
K. Manrique and M. Hermann 1
281
Research on Sweetpotato
Table 1. Mean squares of analysis of variance of AMMI model far total root yield (t/ha) and beta-carotene
content in storage roots (mg/100 g FM) of nine sweetpotato cultivars grown in far locations under two N
regimes (O and 80), Peru, 1999/2000.
BC content
Source of variability
df
Root yield
mg/100 g FM
P>F
t/ha
P>F
2.09
0.0001
Model
87
166.87
0.0001
381.41
0.0001
5.35
0.0001
Enviran
7
1.19
0.0001
Blocks(Ehv)
16
6.0
11.75
0.0001
0.0001
Genotype
8
905.36
0.0001
0.57
0.008
56
80.52
GxE
14
242.79
1.69
PCA1 1
0.38
PCA2 1
12
58.56
0.12
Residual
30
13.57
0.34
Error
128
11.29
Total
215
1 Principal component analysis axes, one and two respectively.
283
PCA2
0.7
XUSHU18
esRON
0.5
WAGABOUGE
0.3
ARB-UNAP 74
DLP2462
TACON
0.1
JPKY 16.005
O.O 1 - - - - - - - - - - - - - - 4 - - - - - - - T - A - C B _ O _ N - - - - - - - - - - - - - - - - 1
-0. 1
eOXAON
eOXA80N
-0.3
LMON
S~2.499-23
-0.5
JEWEL
SR80N e
ARB 535
-0.7 r..---~-~-~-~--~-4---~-~-~--~-~-~-~---.1
-0.6
-0.5
-0.4
-0.3
-0.2
-0.1
o.o
0.1
0.2
0.3
0.4
0.5
0.6
0.7
0.8
PCA 1
Figure 1. Biplot of principal components analysis (PCA) axis 2 vs. axis 1 far beta-carotene content in roots
(mg/100 g) for nine sweetpotato cultivars grown in eight environments.
AMMI biplot PCA 1 vs. PCA2 for betacarotene content in roots for genotypes
and environments (Figure 1) shows stable
cultivars Tanzania, Wagabolige, ARBUNAP 74, Xushu 18, and JPKY 16.005
clustered close to the center of the biplot.
More unstable clones, such as ARB 535,
DLP 2462, SR 92.499-23, and Jewel, were
far from the center. Similarly, environments La Molina (LMON and LM80N) and
Tacna (TACON and TAC80N) are more
stable than Oxapampa (OXAON and
OXA80N) and San Ramon (SRON and
SR80N). This figure also illustrates the
dominant cultivars and environments with
negative PCA 1 seores that strongly influenced the GxE interaction, such as SR
92.499-23, ARB 535, Oxapampa (OXAON
and OXA80N), and San Ramon (SR80N).
3.5
284
Research on Sweetpotato
JPKY 16.005
X JEWEL
XUSHU 18
O SR 92.499-23
.6. WAGABOLIGE
TANZANIA
h. ARBUNAP74
~
u..
8
o
2.5
al
+ ARB 535
1.5
0.5
TAC
LM
SR
OXA
Location
Conclusions
According to the AMMI biplots, none of
the high-yielding cultivars had satisfactory
stability for total root yield. The lack of
association between high root yield and
stable performance suggests the need of
further study on the response of sweetpotato root yield to varying agroecological
conditions. However, promising new highyielding cultivars have been identified. In
Oxapampa, SR 92.499-23 and JPKY 16.005
outperformed the elite cultivar Xushu 18,
which was the top performer at other sites
and has the potential to replace locally
grown varieties in the region.
The biplot for beta-carotene content in
roots showed stability for cultivars Tanzania, Wagabolige, ARB-UNAP 74, Xushu
18, and JPKY 16.005. Clone SR 92.499-23
was less affected by high altitude for betacarotene accumu lation.
There was a trend of increasing betacarotene concentration at increasing
JPKY 16.005
+ ARB535
120.0
DLP2462
TANZANIA
~ ARB-UNAP74
X JEWEL
100.0
XUSHU18
O SR 92.499-23
80.0
.A WAGABOLIGE
60.0
ID
40.0
20.0
O.O
TAC
LM
SR
OXA
Location
Acknowledgments
The authors thank Mr. Adelmo Prraga and
Ms. Pilar Glvez of the Universidad
Nacional Daniel A. Carrin (Cerro de
Paseo) and Dr. Ren Chvez of the
Universidad Nacional Jorge Basadre
(Tacna) for their cooperation in their
respective experimental sites; and field
technicians Mr. Roberto Martinez and
Nstor Crdenas for assisting in plant
harvests and measurements.
285
References
Bacusmo, J.L., W.W. Collins, and A. Jones.
1988. Effects of fertilization on stability
of yield and yield components of sweet
potato. HortScience 113(2):261-264.
Bainbridge, Z., K. Tomlins, K. Wellings,
and A. Westby (eds.). 1996. Methods for
assessing quality characteristics of nongrain starch staples. Part 4. Advanced
methods. Natural Resources lnstitute,
Chatham, U K.
Collins, W., L.G. Wilson, S. Arrende!, and
L.F. Dickey. 1987. Genotype x
environment interactions in sweetpotato
yield and quality factors. Journal of the
American Society of Horticultura!
Science 112(3):579-583.
Gauch, H.G. 1992. Statistical analysis of
regional trials: AMMI analysis of
factorial designs. Elsevier, Amsterdam,
Netherlands. 278 p.
Hammet, H.L. 1974. Total carbohydrate
and carotenoid content of sweet
potatoes as affected by cultivar and
area of production. HortScience
9(5):467-468.
Hussein, M.A. 2000. The statistics of
genotype x environment interaction and
genotype stability in plant breedingtheir computation, statistical test and
interpretation. http://www.nlh.no/ipf/
pu bl i kasjoner/hussei n/stabi lty/
default.htm.
Hussein, M.A., A. Bjornstad, and
A.H. Aastveit. 2000. Sasg x estab, a
SAS program for computi ng genotype x
environment stability statistics.
Agronomy Journal 92:454-459.
Janssens, M.J.J. 1985. Achieving yield
stability in sweetpotato by selection for
translocation potential in adverse
environments. In: Degras, L. (ed.).
Proceeding of the 7th Symposium of the
lnternational Society for Tropical Root
Crops, Gosier, Guadaloupe, 1-6 Jul
1985. lnstitut National de la Recherche
Agronomique (INRA), Paris. France.
p. 729-737.
Jong, S.K. 1974. Genotype x season
interaction and heritability in sweet
286
Research on Sweetpotato
287
289
290
Research on Sweetpotato
Clone selection
The fi rst step in the research reported here
was to analyze the germplasm, consideri ng the ratio of total dry matter of roots to
vines (R/F), and to classify it into five
groups: forage (R/F of 0-1 ), low dualpurpose (R/F > 1-1.5), high dual-purpose
(R/F > 1.5-2.0), low root production (R/F
>2.0-3.0), and high root production (R/F
>3.0) (Len-Velarde et al., 1997). Based on
this classification, 18 accessions were
selected and reclassified into four groups
as follows: group 1, forage (9 accessions);
group 2, low dual-purpose (2 accessions);
group 3, high dual-purpose (2 accessions);
and group 4, low forage-high root production (5 accessions). Accessions classified
as high root production were not included.
Statistical analysis
Four evaluations from years 1999 and
2000, carried out between 120 and 150
days on plots of 1O square meters were
analyzed according to a fixed linear
covariance model:
291
Acknowledgements
This work could not have done without the
fieldwork collaboration of O. Jorges and
C. Aguilar. Many thanks to Z. Huaman,
who facilitated access to all sweetpotato
accessions. Thanks are also deserved by
J. Roca, C. Fonseca, J. Arteaga, L. Quispe,
M. Rojas, C. Gomez, R. Guerra,
References
Arteaga, J. 1997. Potencial forrajero de
cuatro clones nativos de camote
(lpomea batata [L] Lam) para la
alimentacin animal en Oxapampa.
Master's thesis. Universidad Nacional
Daniel Alcides Carrin, Oxapamapa,
Peru. 98 p.
Fonseca, C. 1996. Resultado de las
evaluaciones de un grupo de
colecciones de camote con potencial
forrajero en la Costa Central. lnternational Patato Center, Lima, Peru.
(unpublished data.)
Gomez, C. and E. Quesada. 1 996.
Evaluacin nutricional del silaje de
follaje y raz no comercial de camote
en la alimentacin de vacas lecheras.
Programa de Investigacin en
Alimentos, Facultad de Zootecnia,
Universidad Nacional Agraria La
Malina, Lima, Peru.
Guerra, R. and Len-Velarde. 1998.
Evaluation of sweetpotato and maize.
Interna! document. lnternational Patato
Center, Lima, Peru. (Unpublished data.)
Len-Velarde, CU. 1999. Searching far
dual-purpose sweetpotato in native
germplasm. Interna! document. CIP,
Lima, Peru (Unpublished.) 4 p.
Len-Velarde, C., J. Roca, J. Arteaga,
L. Quispe, and A. Parraga. 1997.
Perpectives on sweet patato: Dual
purpose varieties. In: Program report
1995-1996. lnternational Patato Center,
Lima, Peru. p. 291-294.
Len-Velarde, C. and C. Gomez. 1996. El
camote como forraje animal: Llenando
un vaco en la oferta alimentaria.
293
294
Research on Sweetpotato
295
296
Research on Sweetpotato
Table 1. List of the 113 Latin American sweetpotato accessions used for microsatellite analysis.
Country
Collection
Country
Collection
Country of
Collection
Code
Code
Code
of origin
of origin
No.
origin
No.
No.
1
Peru
CIP ARB 386
39
Colombia
DLP 2104
77
Guatemala
GUA 948
2
DLP 1900
Peru
40
Colombia
DLP 1793
78
Guatemala
GUA STRA
Colombia
DLP 1858
3
Peru
DLP 1922
41
79
Guatemala
GUA 494
4
DLP 206
42
Colombia
DLP 1736
80
Mexico
CTX 15
Peru
43
Colombia
DLP 1771
81
Mexico
CTX7
5
Peru
DLP 2
DLP 1870
82
Mexico
6
Peru
DLP 1090
44
Colombia
CTX 22
DLP 1011
7
Peru
DLP 2344
45
Colombia
83
Mexico
CTX24
DLP 1685
84
Mexico
Peru
ARB 234
46
Colombia
CTX 31
8
DLP 1731
Mexico
DLP 5314
47
Colombia
85
A 160
9
Peru
CTX 33
DLP 3824
48
Colombia
DLP 2046
86
Mexico
10
Peru
11
RCB IN- 90
49
Colombia
DLP 1879
87
Mexico
101438
Peru
Colombia
Mexico
NIAR 221
12
DLP 1921
50
DLP 1737
88
Peru
ARB 355
51
Colombia
DLP 1785
89
Mexico
CATIE 9232
13
Peru
52
Colombia
DLP 2151
90
Mexico
CTX 29
14
Peru
ARB 455
RCB-IF-30
15
Peru
DLP 2298
53
Colombia
DLP 976
91
Mexico
16
Peru
DLP 253
54
Colombia
DLP 971
92
Mexico
CATIE 9257
LL87-1799
17
Peru
RCB IN-199
Colombia
93
Mexico
CTX 31
55
Colombia
DLP 972
94
Mexico
CTX 32
18
Peru
DLP 909
56
DLP 1755
Mexico
19
Ecuador DLP 1192
57
Colombia
95
CTX 12
Venezuela
DLP 2896
Mexico
CTX 34
20
Ecuador DLP 1161
58
96
Venezuela
DLP 869
97
Mexico
CTX5
Ecuador DLP 1456
59
21
Venezuela
DLP 824
98
Mexico
CTX 16
22
Ecuador DLP 1475
60
DLP 806
Mexico
61
Venezuela
99
CTX 9
23
Ecuador DLP 1449
Nicaragua
Venezuela
DLP 868
DLP 4678
24
Ecuador DLP 1447
62
100
DLP 2868
101
Nicaragua
DLP 4686
Ecuador DLP 1156
63
Venezuela
25
Nicaragua
64
Venezuela
DLP 2869
102
DLP 4617
26
Ecuador DLP 1487
Venezuela
DLP 2902
Nicaragua
DLP 4675
Ecuador DLP 1153
65
103
27
Venezuela
DLP 2884
104
Pan ama
DLP 3874
Ecuador DLP 1493
66
28
Venezuela
Pan ama
DLP 3834
67
DLP 2876
105
29
Ecuador DLP 1186
Venezuela
DLP 2838
106
El Salvador SVG 27
Ecuador DLP 1397
68
30
Venezuela
DLP 800
107
El Salvador SVG 12
31
Ecuador DLP 1257
69
Venezuela
DLP 807
El Salvador SVG 24
Ecuador DLP 1149
70
108
32
Venezuela
DLP 822
El Salvador SVG 8
71
109
33
Ecuador DLP 1435
Honduras
72
Venezuela
DLP 792
110
DLP 4545
34
Ecuador DLP 1157
DLP 765
111
Honduras
DLP 4494
73
Venezuela
Ecuador DLP 1231
35
Honduras
DLP 4521
Venezuela
DLP 790
112
74
Ecuador DLP 1498
36
Honduras
DLP 4558
Venezuela
DLP 2881
113
75
37
Ecuador DLP 1484
Venezuela
DLP 791
76
Ecuador DLP 1405
38
Diagnostic, Atlanta, GA, USA). Visualization was accomplished by silver staining
(Bassam et al., 1991 ).
Each SSR variant was treated as an allele.
The alleles were numbered sequentially
starting with the shortest sequence. The
co-run sequence ladder in the gel helped
us to identify single base differences
between any two given samples. The SSR
allele composition of each variety was
determined. We maintained a common
297
Actual heterozygosity
8ecause of the polyploid nature of
sweetpotato, the dosage effects of an SSR
allele (simplex, duplex, triplex, etc.)
cannot be differentiated. Therefore the
allele frequency could not be counted, as
- - 1824217
18242/6
18242/5
18242/4
18242/3
- - - - 18242/2
Figur~
1. SSR-based m~lti-allelic fingerprints in 25 Latin American sweetpotato accessions. Unes and letters on
the nght mark the putat1~e alleles. Allele~ were.numbered starting with the shortest sequence. In this example,
the lowest band (allele) 1s #1 and the h1ghest 1s #7, with alleles 2-6 lying in between these points. When the
autorads were scored, genotypes were scored as 1/0 far each allele.
298
Research on Sweetpotato
Table 2. Regional distribution of allelic diversity in Latin American sweetpotato germplasm. The allelic
diversity is measured by total number of alleles, region-specific alleles, and actual heterozygosity based
on 6 SSR loci.
lndividuals (%)
Region
Total Region- Sample
Dialleles specific size
MonoTriTetraPentaHexaalle les
allelic allelic allelic
allelic
allelic
allelic
Peru-Ecuador
41
47.9
27.4
17.4
o
o
2
38
7.4
Colombia-Venezuela
46
4
29.5
45.3
20.5
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Mesoamerica
37
28.6
42.7
20
8.7
50
8
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14
113
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Total
it can in diploid species, and the expected
heterozygosity (Nei, 1973) cannot be
estimated. However, the observed heterozygosity can be calculated by using the
percentage of multi-allelic accessions. The
actual heterozygosity varies greatly across
the srx loci, ranging from 0.31 (IB255) to
0.92 (IB316), with a mean of 0.60. This
value is reasonable, considering that
sweetpotato is a hexaploid outbreeding
species. The actual heterozygosity in
S. tuberosum was reported to range from
0.477 to 0.502 (Bonierbale et al., 1993).
A similar value (0.495) was found in CIP's
holding of S. andigena using allonym
analysis (Huamn et al., 2000). However,
sweetpotato has a much higher percentage
of tri-al lel ic arid tetra-al lel ic genotypes
than patato does. The present study found
approximately 25% tri- and tetra-allelic
accessions in sweetpotato, but there are
nly 3.2% in patato (Huamn et al.,
2000). In other words, sweetpotato is
general ly mu ch more heterozygous than
patato.
The actual heterozygosity also varies
greatly among the three regions, similar to
allele numbers. lt is 0.714 in Mesoamerica
and 0.705 in Venezuela-Colombia, but it is
only 0.521 in Peru-Ecuador.
The richness and evenness of alleles in the
three tropical American regions are fully
compatible with our recent diversity
analysis for the same three regions using
AFLP (Zhang et al., 2000), which found
that Central America has the highest intraspecific diversity and Peru-Ecuador the
299
Acknowledgements
300
Research on Sweetpotato
References
Austin, D.F. 1988. The taxonomy,
evolution and genetic diversity of
sweetpotatoes and related wild species.
In: Gregory, P. (ed.). Exploration,
maintenance, and utilization of
sweetpotato genetic resources.
lnternational Patato Center, Lima, Peru.
p. 27-60.
Bassam, B.J., G. Caetano-Anolls, and
P.M. Gresshoff. 1991. Fast and sensitive
silver staining of DNA in
polyacrylamide gels. Annals of
Biochemistry 196:80-83.
Bonierbale, M.W., R.I. Plaisted, and
S.D. Tanksley. 1993. A test of the
maximum heterozygosity hypothesis
using molecular markers in tetraploid
potatoes. Theoretical Applied Genetics
86:481-491.
Brown, A.H.D. 1989. Core collection: A
practica! approach to genetic resources
management. Genome 31 :818-824.
Buteler, M.I., R.L. Jarret, and D.R. La
Bonte. 1999. Sequence characterisation
of microsatellites in diploid and
polyploid lpomoea. Theoretical Applied
Genetics 99:123-132.
Buteler, M.I., D.R. La Bonte, and
R.E. Macchiavelli. 1997. Determining
paternity in polyploids: Hexaploid
simulation studies. Euphytica 96:353361.
Doyle, J.J. and J.L. Doyle. 1990. lsolation
of plant DNA from fresh tissue. Focus
12:13-15.
Hu amn, Z. and O.P. Zhang. 1997.
Sweetpotato. In: Fuccillo, D., L. Sears,
and P. Stapleton (eds.). Biodiversity in
100:9-11.
Janes A. 1965. Cytological observations
and ferti 1ity meas u rements of
sweetpotato [/pomoea batatas (L.) Lam].
Proceedings of the American Society of
Horticultura! Science 86:527-537.
Nei, M. 1973. Analysis of gene diversity in
subdivided populations. Proceedings of
the National Academy of Science USA
70:3321-3323.
Nishiyama, l. 1982. Autopolyploidy
evolution of the sweetpotato. In:
Villareal, R.L. and T.D. Griggs (eds.).
Sweet patato, proceedings of first
international symposium. AVRDC Publ.
No. 82-1 72, Tainan, Taiwan, China.
p. 263-274.
Powell, W., G.B. Morgante, C. Andre,
M. Hanafey, J. Vogel, S. Tingey, and
A. Rafal ski. 1 996. The comparison of
2:225-238.
Shiotani, l. and T. Kawase. 1989. Genomic
structure of sweet patato and hexaploids
in lpomoea trfida (H.B.K.) Don. Japan
Journal of Breeding 39:57-66.
Weber, J.L. and P.E May. 1989. Abundant
class of human DNA polymorphism
which can be typed using the
polymerase chain reaction. American
Journal of Human Genetics 44:388-386.
Zhang, O.P., J. Cervantes, Z. Huamn, and
M. Ghislain. 2000. Assessing genetic
diversity of sweet patato (/pomoea
batatas (L.) Lam.) varieties from tropical
America using AFLP. Genetic Resources
and Crop Evolution 47:659-665.
Zhang, O.P., M. Ghislain, Z. Huamn,
A. Golmirzaie. and R.J. Hijmans. 1998.
RAPO variation in sweetpotato
[/pomoea batatas (L.) Lam] varieties
from South America and Papua New
Guinea. Genetic Resources and Crop
Evolution 45:271-277.
Zhang, O.P., Z. Huaman, F. Rodrguez,
and M. Ghislain. 2001. ldentifying
duplicates in sweetpotato [/pomoea'
batatas (L.) Lam] varieties using RAPO.
Acta Horticulturae 546:535-541.
301
303
304
Research on Sweetpotato
Segregation ratio
The assessment of marker dosage was
done by the expected segregation ratios
(presence:absence) of F1 s for AFLP
markers present in one parent in accordance with the allele dosage for three
cytological theories of sweetpotato (Table
1). To classify fragments into dosage
groups, acceptance regions for simplex,
duplex, triplex, and double-simplex
markers were constructed.
Table 1. Expected segregation ratios (presence:absence) far the inheritance of a single gene in a hexaploid
plant, according to three cytological theories based on allele dosage.
Allohexaploid
Marker
Autohexaploid
Tetradiploid
(disomic)
dos e
(hexasomic)
(tetradisomic)
Aa aa aa 1:1
Simplex
Aaaaaa 1:1
Aaaa aa 1:1
aaaa Aa 1:1
AAaa aa 5:1
Aa Aa aa 3:1
Duplex
AAaaaa 4:1
Aaaa Aa 3:1
AA aa aa aaaa AA Aa Aa Aa 7:1
Triplex
AAAaaa 19:1
AAAa aa AAaaAa11:1
AA Aa aa Aaaa AA AAAA aa AA Aa Aa Quadruplex
AAAAaa the 99% confidence level, allowing type 1
error= type 11 error = 0.5%. The null
hypothesis (1 :1 segregation) was tested
against the alternative hypothesis (3:1
segregation) since al 1 non-single dose
ratios would be 3:1 or greater, regardless of
whether sweetpotato is an auto-, autoallo-,
or allopolyploid (Wu et al., 1992).
305
Research on Sweetpotato
Estimation of polyploidy
For species with levels of high polyploidy,
the ratio between the number of detected
repulsion versus coupling linkages may
provide a crude measurement of preferential chromosome pairing (Sorrels, 1992;
Wu et al., 1992). Using Mapmaker's twopoint linkage analysis (LOO = 4, r < 0.3),
the repulsion linkage was analyzed
between an original marker and inverted
markers.
Segregation ratio
Clearly scorable markers were separated
according to their presence in the male
and female parents and in both. The
observed segregation ratio for, each of the
808 female and 641 male markers are
shown in Figures 1 and 2. The distributions
appear with a large group centered around
35
30
25
>.
o
e
(])
20
:J
CT
(])
u:
15
10
5
o
30
40
35
45
50
55
60
65
70
75
80
85
90
95
100
40
35
30
>.
25
o
e
(])
:J
CT
(])
u:
20
15
10
10
19
28
37
46
55
64
73
82
91
100
307
308
Research on Sweetpotato
(QTL). The percentages of mapped simplex, duplex, and triplex markers were
90%, 54%, and 30%, respectively.
Genome coverage
Acknowledgements
We wish to thank G. Rossel,
D. Carbajulca, and O. Hurtado far their
assistance in lab work. We also thank
J. Schmidt and M. Ghislain far valuable
discussion of the manuscript.
References
Alonso-Blanco, C., A.J.M. Peters,
M. Koorneef, C. Lister, C. Dean, N. van
den Bosh, J. Pot, and M.T.R. Kuiper.
1998. Oevelopment of an AFLP based
linkage map of Ler, Col and Cvi
Arabidopsis thaliana ecotypes and
construction of a Ler/Cvi recombinant
inbred line population. Plant Journal
14:259-271.
Bishop, O.T., C. Cannings, M. Skolnick,
and J.A. Williamson. 1983. The number
309
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31 O
Research on Sweetpotato
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of polymorphic clones required to map
the human genome. In: Weir, B.S. (ed.).
Statistical analysis of DNA sequence
data, Marcel Dekker, NY, USA.
p. 181-200.
Da Silva, J.A.G. 1993. A methodology for
genome mapping of autopolyploids and
its application to sugarcane (Saccharum
spp.). PhD thesis. Cornell University,
lthaca, NY, USA.
Da Silva, J.A.G. and M.E. Sorells. 1996.
Linkage analysis in polyploids using
molecular markers. In: Jauhar, P. (ed.).
Methods of genome analysis in plants:
Their merits and pitfalls. CRC Press,
Boca Raton, FL, USA.
Doyle J.J. and J.L. Doyle. 1990. lsolation
of plant DNA from fresh tissue. Focus
12:13-15.
Grattapaglia, D. and R. Sederoff. 1994.
Genetic linkage maps of Eucalyptus
grandis and Eucalyptus urophylla
using a pseudo-testcross: Mapping
strategy and RAPO markers. Genetics
137:1121-1137.
Jones, A. 1965. Cytological observations
and ferti 1ity meas u rements of
sweetpotato [/pomoea batatas (L.) Lam].
312
Research on Sweetpotato
313
Although originally domesticated in tropical America, the sweetpotato (lpomoea batatas (L.) Lam.) has a long cultivation history in Oceania. Whjle the
post-Columbus dispersal of sweetpotato to Asia and Oceania is well documented, the hypothesis that there was prehistoric transfer by Peruvian or
Polynesian voyagers from Peru to Oceania has long been a controversial
issue. The objective of this study was to assess the genetic diversity and
interrelationship of sweetpotato cultivars from Oceania and Latin America
and to test the hypothesis of human transfer of this crop to the Pacific lslands
in prehistoric time. Seventy-six sweetpotato cultivars from Peru-Ecuador,
Mexico, the Philippines and eight Oceania countries were analyzed using
amplified fragment length polymorphism (AFLP). Multidimensional scaling
(MDS) and analysis of molecular variance (AMOVA) revealed wide genetic
variation in the Oceania gene pool, greater than that of Peru-Ecuador. There
was a significant sweetpotato "gene flow" from Mexico to Oceania. In
contrast, there is little association between the Peru-Ecuador germplasm and
that of Oceania. These results suggest that Peru-Ecuador may not be the
source of the Oceania germplasm. Natural dispersal from Mesoamerica is an
alternative explanation to the 'Kumara hypothesis' for the origin of the
Oceania sweetpotato.
Sweetpotato (/pomoea batatas (L.) Lam.)
was originally domesticated in tropical
America (Austin, 1988; Yen, 1982). The
exact center of origin and domestication
of the sweetpotato has not been wel 1
defined, neither has the wild ancestor of
this species been found. Based on the
numerical analysis of key morphological
characters of sweetpotato and the wild
lpomoea species, Austin (1988) postulated
that sweetpotato originated in the region
between the Yucatn Peninsula of Mexico
and the Orinoco River in Venezuela,
within which the four major American
1
315
316
Research on Sweetpotato
Table 1. CIP number and country of origin of 76 cultivars from Oceania, the Philippines, and Latin America.
CIP No.
Country
CIP No.
Country
CIP 441124
Solomon lslands
CIP 440399
New Caledonia
CIP441126
Solomon lslands
CIP 440294
Cook lslands
CIP 441125
Solomon lslands
CIP 440447
Cook lslands
CIP 441119
Solomon lslands
CIP 440454
Fiji
Fiji
CIP 441116
Solomon lslands
CIP 440456
Solomon lslands
CIP ARB 386
CIP 441120
Peru
ARB 234
Peru
CIP 441117
Solomon lslands
CIP 441123
Solomon lslands
DLP 1900
Peru
DLP 5314
Peru
CIP 441127
Solomon lslands
Peru
Solomon lslands
DLP1922
CIP 441118
Peru
DLP 3824
CIP 441221
Tonga
DLP 206
Peru
CIP 440276
Tonga
DLP 2
Tonga
CIP 441222
Peru
DLP 1090
Tonga
Peru
CIP 440273
Peru
CIP 440274
DLP 2344
Tonga
EECH 18
Peru
CIP 440272
Tonga
Tonga
DLP 1188
CIP 440277
Ecuador
Papua New Guinea
DLP 1161
Ecuador
CIP 440693
Ecuador
Papua New Guinea
DLP 1484
CIP 440129
DLP 1192
CIP 440706
Papua New Guinea
Ecuador
DLP 1449
Ecuador
CIP 440695
Papua New Guinea
DLP 1447
CIP 440696
Papua New Guinea
Ecuador
Papua New Guinea
DLP 1487
Ecuador
CIP 440699
Ecuador
DLP 1156
CIP 440297
Papua New Guinea
DLP 1456
Ecuador
CIP 440296
Papua New Guinea
Ecuador
Papua New Guinea
DLP 1403
CIP 440130
DLP 1423
Philippines
Ecuador
CIP 440660
NIAR 221
Mexico
Philippines
CIP 440657
Philippines
CTX 15
CIP 440290
Mexico
CIP 440669
CATIE 9360
Mexico
Philippines
Philippines
Mexico
CIP 440683
CATIE 9232
Philippines
Mexico
CIP 440667
CTX 29
RCB-IF-30
CIP 440671
Mexico
Philippines
Philippines
CATIE 9257
Mexico
CIP 440666
Mexico
Philippines
CTX 33
CIP 440659
Mexico
Philippines
CTX 31
CIP 440665
Mexico
Philippines
101438
CIP 440676
Mexico
Philippines
CTX 32
CIP 440681
of EcoRl/Msel. After the 1igation of the
ol igonucleotide adapters, the restrictionDNA fragments were amplified using a
polymerase chain reaction (PCR). Primer
annealing is targeted at the adapter and
restriction-site sequence. Three-nucleotide
extensions on both the EcoRI and Msel
primers cause selective ampl ification of
fragments. Primer combinations were
chosen based on the numbers of polymorphic fragments in a set of 1O sweetpotato
genotypes.
317
70
60
50
40
30
Results
Polymorphic bands and multidimensional
scaling
The 1O primer combinations generated 21 O
polymorphic, clearly scorable fragments
for the 76 cultivars. The number of polymorphic bands for ali the Oceania
countries were comparable to those found
in the Mexican accessions and higher than
those of Peru-Ecuador, except for Papua
New Guinea. Moreover, there were 18
bands (9% of the total scored bands) that
were present only in the Oceania and
Mexican accessions.
The MDS plot (Figure 1) showed a weak
relationship between accession and
geographic origin, except for Peru-Ecuador, where the cu ltivars were clearly
distinguishable from the rest. Ali of the 22
*
!
*
* *. .,.,:.* * !
! !
*
!
*
20
10
o
-10
-20
-30
-40
-50
Oceania
-60
-70
- 70
- 60
- 50
- 40
- 30
- 20
- 10
10
20
30
40
50
60
70
Figure 1. lnterrelationship of sweetpotato cultivars from America and Oceania using multi-dimensional scaling
(MDS). There is clear differentiation between Peru-Ecuador and the other countries. Mexican cultivars are closely
associated with those of Oceania.
318
Research on Sweetpotato
Table 2. Analysis of molecular variance far the extraction of components of AFLP variation among
regions, and among individuals within regions.
% Total 3
Source of variation
DF
MSD2
Variance
SSD1
component
Among groups (Peru-Ecuador vs. the other
201.2 201.2
4.97
12.2
regions)
Regions within group (among Mexico, the
42.0
0.547
2
84.1
1.4
Philippines, and Oceania)
lndividuals within regions (or countries)
72
2486.5
35.0
35.0
86.4
31.5
Peru-Ecuador
21
661.8
337.3
33.7
Mexico
10
407.1
37.0
Philippines
11
Oceania
276.9
36.0
30
Total
75
groups,
P value4
<.001
NS
<.001
3 Percent
319
Discussion
The high level of genetic diversity and the
low within-region geographic proximity in
Oceania agrees with the previous findings
of Jarret and Austin (1994), who found
higher diversity in accessions from
Oceania than in those from Peru, based on
RAPO analysis. The high diversity in Asia
and Oceania was also found by Huang
(2001 ). This high diversity and the low
geographic proximity within Oceania, as
well as between the Oceania countries
and the Philippines and Mexico, are
compatible with the history of the introduction and cultivation of the sweetpotato
in Oceania and Asia. While the original
spread of sweetpotato within Oceania was
largely due to the Maori, who carried it
with them on their voyages, there were
multiple introductions from different
sources after the initial transmission. For
example, there were at least three notable
introductions to New Zealand. There are
three Maori words used to describe
sweetpotatoes, which reflect the introduction of the sweetpotato to New Zealand:
kumara (which, as mentioned above, has
parallels to the Quechua word kuma),
merikana (meaning American), and waina
(meaning vine) (Yen, 1974). In the Philippines, there were introductions via the
trade route between Mexico and Manila in
the 17th century (O'Brien, 1972; Yen, 1982)
in addition to the introduction from China
by the end of the 16th century. Botanical
evidence also suggests introductions from
Indonesia and west New Guinea (Yen,
1991 ). These repeated introductions from
different sources have contributed to the
high diversity in Asia and Oceania, but
they tend to confound any regional
differentiation between these countries.
320
Research on Sweetpotato
References
Armstrong, J., A. Gibbs, R. Peakall, and
G. Weiller. 1995. RAPDistance,
Package Manual. Version 1.03.
Australian National University,
Can berra.
Austin, D.F. 1988. The taxonomy,
evolution and genetic diversity of
sweetpotatoes and related wild species.
In: Gregory, P. (ed.). Exploration,
p. 27-60.
Doyle, J.J. and J.L. Doyle. 1990. lsolation
of plant DNA from fresh tissue. Focus
12:13-15.
Emory, K.P. 1968. Review of reports of the
Norwegian archaeological expedition to
Easter lsland and the east Pacific. Vol.
2: Miscellaneous Papers. American
Anthropologist 70:152-154.
Excoffier, L., P.E. Smouse, and J.M.
Quattro. 1992. Analysis of molecular
variance inferred from metric distances
among DNA haplotypes: Applications to
human mitochondrial DNA restriction
data. Genetics 131 :479-491.
Gichuki, S.T. 2001. Genetic diversity of
sweetpotato (lpomea batatas [L.] Lam.)
as assessed with RAPO markers in
relationship to geographic sources. PhD
thesis. University of Agricultura!
Sciences, Vienna, Austria.
Heyerdahl, T. 1950. The voyage of the raft
Kon-Tiki. Geographic Journal 115:20-41.
Huang, J.C. and M. Sun. 2000. Genetic
diversity and relationships of sweetpotato and its wild relatives in Jpomoea
series Batatas (Convolvulaceae) as
revealed by inter-simple sequence
repeat (ISSR) and restriction analysis of
chloroplast DNA. TAG 100:1050-1060.
Huang, J.C. 2001. Analysis of genetic
diversity in a sweetpotato (Jpomoea
batatas) germplasm collection from Asia
using amplified fragment length
polymorphism (AFLP). PhD thesis. Hong
Kong University, Hong Kong.
Huamn, Z. and O.P. Zhang. 1997.
Sweetpotato. In: D. Fuccillo et al.
(eds.). Biodiversity in Trust.
Conservation and Use of Plant Genetic
Resources in CGIAR Centres.
Cambridge University Press, Cambridge,
UK. p. 29-38.
Jarret, R.L. and D.F. Austin. 1994. Genetic
diversity and systematic relationships in
sweetpotato Upomoea batatas (L.) Lam.)
and related species as revealed by
RAPO analysis. Genetic Resources and
Crop Evolution 41 :165-173.
16(3):1-1 O.
Vos, P., R. Hogers, M. Bleeker, M. Reijans,
T. van der Lee, M. Homes, A. Frijters,
J. Pot, J. Peleman, M. Kuiper, and
M. Zabeau. 1995. AFLP: A new
technique for DNA fingerprinting.
Nucleic Acid Research 23:4407-4414.
Yen, O.E. 1974. The sweetpotato and
Oceania: An essay in ethnobotany.
Bernice P. Bishop Museum Bulletin 236.
Bishop Museum Press, Hawaii, USA.
Yen, O.E. 1982. Sweet potato in historical
perspective. In: Villareal, R.L. and
T.D. Griggs (eds.). Sweet potato,
Proceedings of First lnternational
Symposium. AVRDC Publ. No. 82-172,
Tainan, Taiwan. p. 17-30.
Yen, O.E. 1991. The social impact of
sweetpotato introduction in Asia and the
South Pacific. In: UPWARO sweetpotato
cultures of Asia and South Pacific.
Proceedings of the 2d Annual UPWARO
lntl. Conference. UPWARD, Los Baos,
Philippines. p. 18-27.
Zhang, O.P., M. Ghislain, Z. Huamn,
A. Golmirzaie, and R.J. Hijmans. 1998.
RAPO variation in sweetpotato
[lpomoea batatas (L.) Lam.] cultivars
from South America and Papua New
Guinea. Genetic Resources and Crop
Evolution 45:271-277.
Zhang, O.P., J. Cervantes, Z. Huamn,
E. Carey, and M. Ghislain. 2000.
Assessing genetic diversity of sweet
potato (lpomoea batatas (L.) Lam.)
cultivars from tropical America using
321
UK
323
available. This fraction was then multiplied by the FAO estmate of the average
total national sweetpotato area for 19982000 (FAO, 2001) to create a consistent
database representative for one ti me
period. We used national-level estimates
of total sweetpotato area for the countries
for which FAO did not provide an estmate, namely: Colombia, Costa Rica,
Guatemala, Guyana, and Nepal. For
Malawi we assumed that the 80% of the
total area reported as potato by FAO was
in fact sweetpotato (Peter Ewell, CIP,
1999, pers. comm.)
Data conversion and analysis
324
Research on Sweetpotato
Results
The sweetpotato distribution database has
a total of 1136 spatial units, with a mean
of 8456 and a median of 1569 ha per map
unit. Detailed results for first-level administrative subdivisions (e.g., state,
department) are tabulated in Huaccho and
Hijmans (2000). Figures 1 and 2 are
derived from the database.
The high concentration of sweetpotato
area in China, which has about 65% of the
world's sweetpotato area, is the most
striking aspect of the crop's distribution
(Figure 1). In addition to China, there are
considerable concentrations of
sweetpotato in Cuba and Haiti in the
Caribbean region; in Java (Indonesia), the
island of New Guinea (both in Indonesia
and in Papua New Guinea), and Vietnam
in Asia; and in Africa, particularly in the
Lake Victoria area (Burundi, Rwanda,
Uganda, and the Dem. Rep. of Congo),
and in Ghana, Nigeria, and Madagascar.
Sweetpotato is an important staple for
many Oceana island countries such as
Papua New Guinea, the Solomon lslands,
Tonga, and New Caledonia. They are not
shown on the map, however, because of
the 1-dot-equals-1 000-ha scale used.
Because of the high concentration of
sweetpotato area in China, we include a
separate map for that country (Figure 2). In
China, there are three areas of very high
concentration of sweetpotato: 1) Sichuan
and Chongq i ng provi nces, with about
1,000,000 ha (the Sichuan Basin); 2) the
east central provinces of Shandong,
Henan, and Anhui, each of which has over
600,000 ha; and 3) the southeastern coast.
Each of four Chinese provinces has more
sweetpotato area than Uganda, which,
with 546,000 ha, is the second largest
sweetpotato producing country.
There is a bimodal distribution of
sweetpotato area by latitude (Figure 3).
Seventy percent of sweetpotato is grown
between 20N and 40N. This peak
includes nearly all the area in China,
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700
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300
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100
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-20
10
-10
20
30
40
50
60
L.atitude()
326
Research on Sweetpotato
Fraction
0.8
0.6
0.4
0.2
50
-40
-30
-20
-10
10
20
30
40
50
60
Latitude ()
50
45
40
35
30
25
20
15
10
45
40
35
30
25
20
15
10
0-5
5-1 o
10-15
15-20
20-25
25-30
Temperatura (C)
327
328
Research on Sweetpotato
Acknowledgments
We thank Terefe Belehu, Peter Ewell, Ali
Khalafalla, Berga Lemaga, Mlance
Ndikumasabo, Jean Ndirigue, Phemba
Phezo, J.M. Randrianaivoarivony, Tom
Walker, and Stanley Wood for providing
sweetpotato distribution data. Additionally,
we thank Tom Walker for reviewing this
paper.
References
Bertin, J., J.J. Hmardinquer, M. Keul, and
W.G.L. Randles. 1971. Atlas of food
crops. Geographical and chronological
survey for an atlas of world history.
cole Pratique des Hautes tudesSorbonne. Vle Section: Sciences
conomiques et Sociales. Centre de
Recherches Historiques et Laboratoire
de Cartographie, Paris, France.
Carter, S.E., L.O. Fresco, and P.G. Janes
with J.N. Fairbairn. 1992. An atlas of
cassava in A frica. H istorical,
agroecological and demographic
aspects of crop distribution. Centro
Internacional de Agricultura Tropical,
Cali, Colombia.
Crook, F.W. 1993. Under reporting of
China's cultivated area: lmplications for
world agricultura! trade. In: China,
international agriculture and trade
reports. Situation and outlook series, RS93-4. United States Department of
Agriculture, Economic Research
Service, Washington, D.C., USA.
De Vries, C.A., J.D. Ferwerda, and M.
Flach. 1967. Choice of food crops in
relation to actual and potential
production in the tropics. Netherlands
Journal of Agricu ltural Science 15: 241248.
FAO (Food and Agriculture Organization
of the United Nations). 2001. Database
at http://apps.fao.org.
Hijmans R.J. 2001. Global distribution of
the patato crop. American Journal of
Patato Research, vol. 78.
329
331
2 PRA
= field observations.
3 Number
332
Research on Sweetpotato
Methods
The CIP/UPWARD monitoring and evaluation study appl ied an analytic framework
developed with all stakeholders during a
workshop at the beginning of the project.
lndicators far impact evaluation had been
determined with the stakeholders earlier
during the ICM research project. The
framework considered the various program
implementation levels far monitoring and
evaluation: 1) training of facilitators,
2) FFS implementation, 3) horizontal
dissemination within the community,
4) ICM implementation by trained farmers,
5) farm level effects, and 6) implications
far research and development institutions.
Far each evaluation leve!, infarmation
sources, variables, and indicators were
determined, and data collection methods
designed as fal lows.
Pre-FFS:
Observations and interna! evaluation
during facilitator training.
Individual qualitative baseline interviews with FFS participants (who later
become ICM farmers).
During FFS:
Process observation and recording
during FFS sessions.
Pre-tests/post-tests on ICM farmers'
knowledge.
Post-FFS:
Evaluation meetings with participants
and faci 1itators.
Individual interviews with facil itators,
village officials, and traders.
Two-step individual interviews with ICM
farmers (FFS graduates) and randomly
selected non-ICM farmers.
333
Results
Due to limitations of space and the fact
that the PRGA analysis is still in process,
only a limited number of key results are
included here. The first set of results
pertains to measurable impacts of the
ICM-FFS on farmers' knowledge, skills,
production practices, and social relations
in the FFS villages. This analysis is
fallowed by a discussion of the role of
farmer participation in the design and
development of the ICM-FFS, and its
implications far impact.
334
Research on Sweetpotato
Table 3. Overview of implementation and process evaluation of six NIPMP-conducted sweetpotato ICM
farmer field schools, 1997-98.
District
Mojokerto
Magetan Karanganyar Magelang
Kuningan
Sleman
Variable
East Java
East Java Central Java Central Java Yogyakarta West Java
yes
Majar sweetpotato
yes
yes
no
no
yes
growing area
FFS implementation
Jan-Jun 98 Aug-Dec 97 Sep 97-Feb Sep-Dec 97 Aug-Nov 97 Nov 97-Apr
season
(weVdry)
98 (wet)
(dry/wet)
(dry/wet)
98 (wet)
(dry/wet)
Meetings (no.)
17
17
17
18
17
17
Participants (range)
10-25
11-27
14-27
11-27
11-27
13-28
Meetings (avg. no.)
16
18
18
19
16
17
Women participation
0%
0%
7%
40%
56%
60%
(% of trainees)
varieties
varieties
Fertilization varieties; N Fertilization
Organic
Experiments
application (potassium,
fertilizer
conducted 1
(application
N-K rates)
rate; organic)
rates
126%
40%
Avg. knowledge
56%
28%
48%
254%
(25)
(10)
(13)
(24)
(19)
(20)
increase between
~re- and ~ost-test2
1 In addition to standard FFS trials.
2 Pre- and post-tests consisted of 10-15 questions each. Seores correspond with the number of correctly answered
questions. Percentage knowledge increase is calculated by ((score post-test- score pre-test)/ score pre- test)* 100%
The figures in the table represent the average knowledge increase of all participants per FFS who did both pre- and
post-test (number of respondents given in parentheses).
335
Mojokerto
Magetan
Karanganyar
Magelang
Sleman
111
lm
FFS plot
Kuningan
10
20
30
40
Sweetpotato storage root yield (t/ha)
50
60
Mojokerto
Magetan
Karanganyar
Magelang
Sleman
Kuningan
o
1o
20
30
40
50
Market price of sweetpotato storage roots
(US$/t)
10
20
30
40
Sweetpotato cultivation cost
(US$/ha)
250
500
750
Net income from sweetpotato
(US$/ha)
50
Mojokerto
Magetan
Karanganyar
Magelang
Sleman
Kuningan
250
500
750
1000
Gross income from sweetpotato
(US$/ha)
Figure 1. Sweetpotato storage root yield, market price, gross and net income, and cultivation costs at six ICM-FFS
sites, by baseline farmers (1994/95 wet season), on the sweetpotato ICM-FFS plot (for seasons, see Table 3), and
by ICM and non-ICM farmers during a post-FFS season. No post-FFS data were available for non-ICM farmers in
Magelang. Note: The high yields obtained on the FFS learning plots likely result not only from application of the ICM
practices, but also from the fact FFS plots are often relatively fertile and intensely managed. Moreover, they are quite
small and extrapolating kg/plot yield to Vha would be expected to overestimate the yield gains.
336
Research on Sweetpotato
Table 4. Differences in knowledge and skills, practices, and inputs and outputs of sweetpotato cultivation
during a post-FFS season by ICM-FFS participation. Magelang is excluded from the analysis because
there are no non-ICM cases in this community.
Variable
Knowledge and skills
Understand the concept natural enemy (%)
Practice routine/thorough field observation (%)
Conduct experiments (%)
Can assess yield within reasonable range (%)
ICM (N=73)
Non-ICM (N =50)
86
52
75/29
16
53
75/47
18
48
Cultural practices
Practice seed selection (%)
Practice good water management (%)
Use N-fertilizer (%/kg N/ha)
Use P-fertilizer (%/kg N/ha)
Use K-fertilizer (%/kg N/ha)
Use organic fertilizer (%/Vha)
Pesticides applications (avg. no./season)
Practice field sanitation after experiencing weevil attack (%)
45
28/20
35/1.3
0.4
49
37
58
92/242
60/58
13/7
29/1.6
0.3
30
107
29
201
20.5
32
391
112
27
237
19.5
30
316
68
96/223
68/78
337
338
Research on Sweetpotato
339
Research on Sweetpotato
Conclusions
Sweetpotato ICM-FFS has contributed to
increased knowledge and skills among
farmers who participated. These human
capital increases have resulted in concrete
changes in production and marketing
practices that contributed to higher yields
and higher income from sweetpotato
production. These results were essentially
achieved by improving the efficiency of
existing practices rather than by introducing totally new technologies. Farmer
participation in the research process,
particularly in the identification of possible areas for improving efficiency,
appears to have contributed to both the
relevance and the impact of the ICM-FFS.
Whereas most of the impacts shown in this
study are significant statistically, they may
seem to be less so in practica! terms. The
magnitude of the differences of individual
parameters between ICM and non-ICM
farmers are often relatively small, but
taken as a whole they indicate the initiation of change in farmers' crop
management behavior. As mentioned
earlier the data may underestimate the
true differences between ICM and nonICM farmers, mainly as a result of the
inability to control for spillover effects
from participants to nonparticipants. Given
that ICM is complex and that increased
knowledge does not immediately result in
changed practices, it is likely that over
time the benefits from ICM-FFS may grow.
Subsequent field visits as part of the PRGA
study gave that impression. A final factor
that could have affected the magnitude of
the observed impacts is the Asian financia!
crisis that hit Indonesia in 1997. The
profitabi 1ity of sweetpotato was drastical ly
reduced, hence the reduced incentives for
farmers to i nvest in it.
Nonetheless, the question of whether the
benefits justify the costs of ICM-FFS is
valid. This issue will be examined in
greater detail in the forthcoming PRGA
study. Participatory development of the
sweetpotato ICM-FFS took several years
References
Asmunati, R., Wiyanto, and E. van de
Fliert. 1999. lntegrating sweetpotato
ICM and FFS approaches in government
and NGO initiatives in Indonesia, In:
Users' Perspectives for Agricultura!
Research and Development (UPWARD),
Learning to manage livelihoods: New
perspectives in rootcrop R&D.
341
342
Research on Sweetpotato
343
Data are reported only for the six provinces considered key sweetpotato
production areas. Hoa Binh and Dong Nai
provinces, which were studied only for
canna production, are not included in the
analysis.
Methods
The study emphasized two key types of
baseline information, i.e., 1) production
and crop management, and 2) postharvest
use in both sweetpotato and canna systems. Study sites represented key
sweetpotato or canna production areas,
and included eight provinces across the
country.
Northern Vietnam: Vinh Phuc and Ha
Bac provinces (sweetpotato), and Hoa
Binh (canna).
Central Vietnam: Thanh Hoa Province
(sweetpotato and canna), and Quang
Nam Province (sweetpotato).
Southern Vietnam: Ba Ria-Vung Tau and
Vinh Long provinces (sweetpotato), and
Dong Nai Province (canna).
344
Research on Sweetpotato
Results
Characterization of sweetpotato
cultivation areas and systems
Sweetpotato cultivation in the various
regions in Vietnam varies by agroecology
and climate. In all study sites, sweetpotato
is typically grown as a field crop in
rotation with rice and other crops such as
maize, groundnut, cassava, and vegetables. Winter and spring constitute the
major cultivation season in northern and
central Vietnam. In southern Vietnam
sweetpotato is grown year-round, but
preferably during the dry season, which
coincides with the winter-spring season in
the North. Women have a major role in
sweetpotato cu ltivation, particularly in
northern and north-central Vietnam.
Further to the south the average proportion
of work done by women is smal ler (Table
1), which is consistent with results of
Tuyen (1999). Sweetpotato ranks first
among major field crops in Ha Bac, Thanh
Hoa (one district), Ba Ria-Vung Tau, and
Vinh Long provinces, and either second,
third, or fourth (depending on a specific
community) in Vinh Phuc, Quang Nam,
and Thanh Hoa. Sweetpotato is favored
because of its high productivity and low
management and input requirements,
which makes it an easy and potentially
profitable enterprise. Additional ly, seed is
readily available, and in sorne areas
sweetpotato is one of the few crops
Sweetpotato production
In all study sites, farmers plant sweetpotato on ridges, although the width and
height, and accordingly plant population,
vary by soil type, water supply, and local
CIP Program Report 1999- 2000
345
w
.:..
O)
o
:::1
en
$.
'O
Table 1. Sweetpotato (SP) production and postharvest use characteristics in six needs assessment sites in Vietnam, 1998/99 .
Northern Vietnam
Central Vietnam
Characteristic
Southern Vietnam
Vinh Phuc
Ha Bac
Thanh Hoa
Quang Nam
Ba Ria-Vung Tau
Vinh Long
1:0.3
1:0.4
1:0.6
1:1.1
1:1.6
Labor ratio,
1:1.4
women:men (h/ha)
1, Befare rice,
1, Befare rice, beans,
2, 3, or 4, After
Rank of SP among
3, 4, After rice,
1, Befare rice
1, Befare rice,
maize, groundnut groundnut. 2, 3, after
other majar crops
maize, sugarcane,
cassava, rice,
maize, vegetables
groundnut, rice
groundnut
mulberry, soybean
Typical cultivation
pattern 1
Rice - SP or
vegetables - rice
or groundnut
Rice - SP Vegetables or
cassava/ sesame
Rice - SP
Most serious
production
constraints
SP weevil
Low soil fertility
Lack of suitable
varieties
Soil fertility
SP stem borer
Virus es
Drought, cold
Lack of suitable
varieties
Lack of capital
SP weevil
Lack of capital
Heavy rain
Soil fertility
Fairly fertile
Moderately fertile
Fairl~ infertile
Water availability
Fairly reliable
Moderately reliable
Fairly unreliable
1 Spaced
38%
50%
13%
63%
38%
0%
73%
28%
0%
3%
53%
45%
100%
0%
0%
0%
25%
75%
58%
43%
0%
0%
3%
98%
23%
15%
63%
0%
0%
100%
100%
0%
0%
hyphen indicates followed by. For example: rice crop followed by a sweetpotato crop indicated as rice - sweetpotato.
Table 2. Contribution to gross income of the various uses of vines and storage roots, as average percent of
total gross income (including opportunity value far in-kind products) from sweetpotato. Vietnam, 1998/99.
Use
Northern Vietnam
Central Vietnam
Southern Vietnam
Vinh Phuc
Ha Bac
Thanh Hoa Quang Nam
Ba Ria-V. Tau Vinh Long
{n=40}
{n=40}
{n=40}
{n=40}
{n=25}
{n=40}
Vines:
38.5
31.5
62.5
29.6
O.O
0.2
o.o
Animal feed
24.0
18.7
29.7
23.2
o.o
12.7
28.6
4.2
o.o
Sold to market
10.7
0.2
4.2
2.1
Seed
2.9
O.O
O.O
O.O
0.8
O.O
O.O
0.1
O.O
Give away/
O.O
other uses
37.5
70.4
100.0
Storage roots:
61.5
68.5
99.8
99.3
97.5
11.3
Sold to market
10.2
39.4
13.6
17.0
0.1
7.8
0.1
21.3
Animal feed
25.4
19.1
0.1
Family
5.8
5.8
1.6
13.0
consumption
O.O
O.O
O.O
10.4
22.9
Used far
1.0
processing
O.O
O.O
2.0
0.1
O.O
Discarded 1
3.9
O.O
O.O
O.O
0.5
0.6
Give away/
8.1
other uses
1 Due to weevils, rot and other causes.
34 7
Table 3. Sweetpotata production practices, farmer knowledge and skills in the six needs
(% farmers reQorting unless SQecified otherwise).
Variable
Northern Vietnam Central Vietnam
Ha
Vinh
Thanh Quang
Bac
Phuc
Hoa
Nam
{n=40} {n=40} {n=40} {n=40}
Sail preparatian and planting:
Avg. width af ridge (cm)
80
67
107
199
Avg. height of ridge (cm)
40
34
54
58
Avg. planting density ('000 plants/ha)
67
48
47
54
Practice seed selectian
98
100
98
100
Apply organic manure
8
100
100
100
Apply fertilizer
lnorganic N
95
100
100
100
lnorganic P
o
13
95
8
lnarganic K
o
88
100
33
Apply pesticides
o
o
o
85
Practice field manitoring:
Occasionally
o
18
80
98
Routinely
100
83
20
3
Understand the concept of natural enemies
o
13
8
70
Can assess root yield within reasonable range
45
35
15
38
Market prices
Far vines (01/kg) (avg.)
348
216
400
505
Far roats (D/kg) (avg.)
624
534
716
503
Stare sweetpotato roats
85
100
43
100
assessment sites
Southern Vietnam
Ba RiaVinh
Vung Tau Long
{n=25} {n=40}
133
70
64
100
100
100
100
100
57
40
68
100
98
48
53
98
3
3
100
100
28
1100
18
1000
1435
348
Research on Sweetpotato
1,600
40
Vines
Roots
1,400
1,200
30
1,000
en
2
800
?
.e
2!. 20
Labor
~ lnputs
Net return
(])
"'O
Q5
>=
?
.e
600
10
-~
400
200
en
en
C)
Province
Province
Figure 1: (A) Average total yield broken down into average storage root yield and vine yield (left). (B) Average
gross income broken down into net income and cost for inputs and labor.
Table 4. Multiple linear regression of cultivation variables possibly accounting for sweetpotato vine and
storage root yield (dependent variable). lndependent variables include plant density, labor, crop growth
duration, doses of organic manure and inorganic (N, P, K, and others which is mainly NPK composite)
fertilizers, and pesticide application frequency. Yalues per entry are standardized coefficient and probability
P (in parenthesis). Only values far independent variables with a probability of P<0.1 O are displayed.
lndependent variables
Adjusted Density
Growth
Labor
Fertilizer
Pesticide
R2
(plants/ha) duration
(hr/ha)
N (kg/ha) K (kg/ha) Organic
application
(days)
(kg/ha)
frequency
Vines
0.514
Vinh Phuc
0.225
(0.006)
-0.354
0.365
0.198
Ha Bac
(0.068)
(0.059)
0.303
Thanh Hoa 0.032
(0.094)
-0.914
0.223
0.239
Quang Nam 0.773
(0.022)
(0.013)
(<0.001)
Roots
0.162
0.389
Vinh Phuc
(0.040)
0.440
-0.538
0.329
0.213
Ha Bac
(0.009)
(0.086)
(0.008)
0.383
Thanh Hoa 0.015
(0.038)
0.007
0.188
Quang Nam 0.816
-0.883
(0.240)
(0.028)
(<0.001)
-0.043
Ba RiaVung Tau
-0.122
0.301
0.773
Vinh Long
0.869
(0.062)
(<0.001)
(0.018)
349
Research on Sweetpotato
Discussion
Sweetpotato productivity in Vietnam is
general ly low. But in the study sites, where
one would expect the crop to be well
suited to local conditions, average storage
root yields were higher than the national
average of 6.5 t/ha (FAO, 2000). Low
Conclusions
The i ntegrated needs assessment reported
here is the most comprehensive study done
to date in Vietnam on root crop production-postharvest use systems. Containing as
it does production, post-production, and
socioeconomic elements, it provides a
fairly complete picture of the constraints
References
Bottema, J.W.T., P.T. Binh, D.T. Ha,
M.T. Hoanh, and H. Kim. 1991. Sweet
potato in Viet Nam, production and
markets. CGPRT No. 24. CGPRT, Bogor,
Indonesia. 113 p.
FAO. 2000. FAOSTAT Database, http://
apps. fao.org/.
Tuyen, N.N. 1999. Women in Vietnam's
national IPM program, In: van de Fliert,
E. and J. Proost. Women and IPM: Crop
protection practices and strategies. KIT
Press, Royal Tropical lnstitute,
Amsterdam, Netherlands. p. 79-88.
Van de Fliert, E. and A.R. Braun. 1999.
Farmer field school for integrated crop
management of sweetpotato: Field
guides and technical manual.
lnternational Potato Center, Bogor,
Indonesia. 286 p.
Walker, T. and M.H. Collion. 1997. Priority
setting at CIP for the 1998-2000
Medium-Term Plan. lnternational Potato
Center, Lima, Peru. 48 p.
351
352
Research on Sweetpotato
355
Table 1. General description of the selected hydrologic units showing altitude, predominant soil type,
average soil organic matter (OM) content, total area cropped to patato, and amount of the insecticide
carbofuran applied to patato fields.
Soil type 1
Hydrologic
Altitude (m)
OM content
Total area
Patato area
Carbofuran
(%)
(ha)
(ha)
applied 2
unit number
(l/ha)
Cf
7.0
2.0
2.00
1
3070
8
Dp
10
7.0
6.0
2.00
2
3200
Dp
5.0
2.2
0.00
3100
10
3
Dp
10
20.0
4.0
2.00
4
3100
Dp
10
10.0
4.0
4.00
5
3000
Dp
12.0
5.0
3100
10
2.00
6
Dp
10
2.5
1.5
0.00
7
3100
Dp
10
6.0
1.2
2.30
3200
8
Cf
8
10.0
5.0
2.00
9
2900
Dp
8.0
7.0
10
3100
10
2.00
Hf
6
3.0
3.0
11
2770
0.25
12
Hf
6
7.0
6.5
2900
1.60
13
2800
Hf
6
4.5
4.0
4.50
4.0
14
2900
Cf
8
1.5
1.50
1 Soil types from MAG-ORSTOM (1980). "Cf" stands for Duriudoll-Eutrandept, "Dp" for Dystrandept and "Hf" for
Eutrandept-Argiudoll.
2 Amount of commercial formulation (Carbodan, Carbofuran 4f) applied during potato cycle.
356
Top position
Lateral
movement
of water
Surface water
(creek or ditch)
Figure 1. Representation of a sloping field and the relative positions (top, middle, and bottom) from which soil
leachate, groundwater, and surface water samples were taken.
a depth of 80 cm just below potato roots. lf
a sandy or cemented layer was present at
a shallower depth, the probes were placed
above that layer. We were able to extract
sufficient soil solution for carbofuran
analysis during a 16-24 h period. Soil
leachate samples were taken in this way
from a total of 19 fields across ali hydrologic units.
An attempt to collect groundwater samples
was made once during the sampling period
by boring a hole to a depth of 220 cm at
the bottom portian of one field within each
hydrologic unit. lf groundwater was found,
it was carefully pumped out using a
syringe attached to a plastic pipe; in only
7 of the 14 units did we encounter groundwater at a depth of 220 cm or less. Surface
water samples were taken usual ly at least
twice, once near the beginning of the
sampling period and again near the end,
by scooping plastic bottles into water
running through a creek or ditch at the
bottom of each hydrologic unit.
Leachate and water samples were analyzed for the presence of carbofuran using
the Carbofuran RaPID Assay procedure
(Strategic Diagnostic, lnc., http://
Model simulations
To simulate carbofuran movement in soil,
we used the pesticide version (LEACHP) of
the basic LEACHM model (Hutson and
Wagenet, 1992). Simulation runs were
made for each potato field sampled in the
14 hydrologic units. To run the model,
weather data inputs (rainfall and temperature) were obtained from the INAMHI
(Instituto Nacional de Meteorologa e
Hidrologa) meteorological station in
nearby San Gabriel, although for two of
the units rainfall was measured on site.
Soil data inputs (physical and chemical
properties) were estimated in the field
according to functional horizon relationships derived earlier by Kooistra and
Meyles (1997). Carbofuran amounts and
application dates for each field were
obtained from farmer interviews.
357
Table 2. Carbofuran concentrations (parts per billion (ppb)) found in soil leachate, ground water, and
surface water in the fourteen hydrologic units.
Carbofuran in soil leachate
Hydrologic
Soil type
Carbofuran
Carbofuran
Mean
Maximum
Minimum
in ground
in surface
unit number
(ppb)
(ppb)
(ppb)
water1(ppb) water12 (ppb)
Cf
1.00
4.00
0.00
ns
0.10
1
Dp
0.00
0.00
0.19; 0.00
0.00
0.00
2
Dp
0.12;0.00
0.04
0.10
0.00
ns
3
Dp
0.02
0.12
0.00
0.15
0.00
4
Dp
0.26
0.52
0.00; 0.65
0.10
ns
5
Dp
0.00
0.00
0.00
0.00; 1.60
6
ns
Dp
0.00
0.00
0.00
0.00; 2.00
7
ns
Dp
0.63
2.50
8
0.00
0.00
0.00; 0.07
Cf
0.05
0.20
0.00; 0.95
9
0.00
ns
Dp
0.00
0.00
0.00
0.00
ns
10
1.28
11
Hf
1.80
0.27; 0.80
0.58
0.36
Hf
1.82
6.20
0.29; 0.58
12
0.40
0.32
Hf
1.22
13
3.50
0.00
0.11; 0.26
0.75
14
Cf
1.23
3.60
0.00
ns
0.50; 0.46
1
ns = no sample available.
Surface water samples were taken once or twice. Where two samples were taken, the first number is the first sample
result; the second number is the second sample result.
358
Conclusions
Carbofuran was found in soil leachate
groundwater, and surface water sampl~s
taken from sites within 14 hydrologic units
in Carchi. Analysis confirmed a low
degree of pollution for the area that is
below the MCL of 40 ppb, but often above
a level of 0.4 ppb known to adversely
affect sorne aquatic 1ife forms. Des pite the
high OM content of the soils, and evidence for the fairly rapid degradation of
carbofuran in these environments, leaching of carbofuran does occur and it can
make its way into groundwater.
Model simulations
The simulation of carbofuran movement
with LEACHM showed the model to be
extremely sensitive to the degradation
rate, or half-life used for carbofuran. When
the more conservative estimate of pesticide persistence was used (half-life = 50
d), the model predictions of carbofuran in
soil leachate greatly exceeded the amount
measured (Figure 2). Nevertheless, the
simulations did capture the inverse
relationship between soil OM content and
the amount of carbofuran leached. When
the simulation used the half-life of 14 d
that was obtained from the experiment
conducted local ly by Stoorvogel et al.
(2001 ), model predictions were closer to
:e
.t:
W Measured carbofuran
JI
Simulated carbofuran
!,,= 50 days
O Simulated carbofuran
t,.= 14 days
CJ
al
e
e
al
Hf(6%SOM)
Cf(8%SOM)
Dp(10%SOM)
Soiltypes
359
References
Crepeau, K.L. and K.M. Kuivila. 2000.
Rice pesticide concentrations in the
Colusa Basin Drain and the Sacramento
River, California, 1990-1993. Journal of
Environmental Quality 29:926-935.
Crissman, C.C., J.M. Antle, and S.M.
Capalbo (eds.). 1998a. Economic
environmental and health tradeotfs in
agriculture: Pesticides and the
sustainability of Andean patato
production. Kluwer, Boston, MA, USA.
275 p.
Crissman, C.C., P. Espinosa, C.E.H.
Ducrot, D.C. Cole, and F. Carpio.
1998b. The case study site: Physical,
health and patato farming systems in
Carchi Province. In: Crissman, C.C.,
J.M. Antle, and S.M. Capalbo (eds.).
1998. Economic, environmental and
health tradeoffs in agriculture:
Pesticides and the sustainability of
Andean patato production. Kluwer,
Boston, MA, USA. p. 85-120.
Ducrot, C.E.H., J.L. Hutson, and R.J.
Wagenet. 1998. Describing pesticide
movement in patato production on
Carchi soils. In: Crissman, C.C., J.M.
Antle, and S.M. Capalbo (eds.). 1998.
Economic, environmental and health
tradeoffs in agriculture: Pesticides and
the sustainability of Andean patato
production. Kluwer, Boston, MA, USA.
p. 181-208.
EPA (Environmental Protection Agency).
1998. Drinking water and health.
Available at Environmental Protection
360
1
2
Location
The llave-Huenque watershed (38255550 m) of the Andean high plateau or
Altiplano was the subject of this case
study. This is one of the most important
watersheds; its effluent drains into Lake
Titicaca, sustaining the lives of thousands
of resource-poor households that depend
on agriculture. The northernmost points in
361
Photogrammetric
charts
Rain
(GOES)
Contour
lines
Tmax,Tmin
Rainfall
(Weather station)
IDRISI
CU MATE
INTERPOLAR
Tmax,Tmin
Rainfall maps
Landsat-TM
IDRISI
NOAAAVHRR
ENVIJIDRISI
ENVI
,,
Soil maps
Climae
index
,,
Land
use/cover
Biomass
,,
,,
ENVI
AE Zones
Figure 1. Flow Diagram of the method for agroecological zoning (Tmax = maximum temperature, Tmin = minimum
temperature). Abbreviations: ARC/INFO software= (www.esri.com/software), AVHRR = Advanced Very High
Resolution Radiometer (NOAA), DEM = digital elevation model, ENVI = Environment for Visualizing lmages
(Research Systems lnc.-Kodak, Boulder, CO, USA), GOES = Geostationary Operational Environmental Satellite
(NOAA, USA), IDRISI = IDRISI software (http://www.clarklabs.org/), NOAA = National Oceanic and
Atmospheric Administration (USA).
the watersheds are at latitude 161 O' S
and longitude 6930' W and southernmost at latitude 1705' S and longitude
7005' W. The total area comprises about
777,000 ha.
362
Soils
Thirty-three soil classes were used in the
AEZ. In general terms, the soils are mostly
shallow to moderately shallow, low in
organic matter content (less than 4%), and
1 318
R1 =
SStotal
pp + 1
R2=0.90
ltr
[ ((T
=7.1 + Ln
- T.)+ 1)1.66
max
mm
363
Soil series
......
SoH t t rl o
c::J
...
]
c:;:;J
Caboono
Cal.3collo Aftor1mit nto s rtioos
CaliJ00Jlt,.C1o hla
C31aoollo Huaitirt:
MI pals
Materia1ts tutaceos
Cilaoollo -M :iloomayo
Csllco lloMltrl:tlu tut>u os
- Pok e
~
~
PoktMaleomayo
Poke Yanam1110
Yanuoo haAlpa (suptricial
c::J Y.tnamayo -1flor.1mltntof ~t loot
Yanamayo -Malcomayo
C31acollo - Poke
c:J
~
c:J
Cabcollo Such u
Cabtoo No To rrtni
Cal3collo -\'tz oaohas
~ Jc o llo
Yanamiyo
Chlno he ros
~iric heros- Ator3miento
Chinchuos lffpa
Chinohtros LIJbo
Cul anel lpa
Hu:altrt: -'Vtzoa chas
Centros pobl;idos
Lagu nu
- H i val
hico
Cllinchtro s Ch tjemooo
Slopes
......
...
,..
...'.
""''"
11
Altitude
......
m.s .n.m .
o~
c:::J
,...,.'
..
,.. '
.,.,
...
'
......
<38 00
3'00 . 31'00
3900. 4000
400 0. 4100
4100 . 4200
4100. 4300
4300 . 4400
4400 4600
4500. 41<!0
...
......
,..
04
4 . 15
16 . 26
25 . 50
>60
......
* . 4700
4900
~
~
470 0
4800 4llOO
4QOO 5000
6000 . 6100
5100. 5200
5200 . 5300
5300 . 6400
Klom t ttfS
20
40
,. $400
'
365
1- -
...
.,.
......
C1tegory
J!
Gt1ssland
CJ
8
Pttt:nnbl s nowhlds
!
B
,..
...
Thennal - ra in lndex
!!
.......
...... ..11
8
8
8
8
...
ti
,..
...
e!l
,..
1B
Biomass
......
lndex
-2
CJO
E3!
.. CJ5
E3~
CJS
. ': ~
~t '
...
,..
'
" :ro;!:
- ~ 10
~~
..._
......
Kg/H 4.
'1000
1000 . 2000
2000. 3000
CJ3n00 . 4000
'4000 . 6000
6000 . eooo
1
,..
"
) 1000.
...
.......
K$omt: ttrS
20
40
7000
......
Figure 3. Land cover and land use, biomass, and climate of llave-Huenque watershed.
Biomass
Us in g N DVI to estimate stand ing green
biomass proved to be a reliable so urce of
biomass data. There are two sources of
data that co mp lement eac h other. On the
one hand, high-reso luti on biomass maps
ca n be deri ved from Landsat-TM (30- m
366
reso lut ion) or from lko nos (4-m reso luti on).
Th e tradeoff usi ng these data is the hi gher
cost. O n th e other hand, lower reso lution
(1 -km) AVH RR might be used. The
advantage of th is se nsor is its co nt inu ous
and synopti c coverage plu s the
ava il ability of data o n the Internet (USGSEOAAC, 2001).
AEZ
Four AEZs were derived from the analysis
(Figure 4). The zone with aptitude for crop
and pasture production comprises 42,000
ha. Using process-based models to
simulate the potential production of this
zone indicates that productivity can be
significantly increased with technologies
to intensify agriculture in the zone. For
example, patato production could be
i ncreased from 5-6 t/ha to 10-12 t/ha
under rainfed cond itions; up to 18 t/ha
with i rrigation.
The second zone corresponds to the area
where livestock can be intensified. lt
comprises roughly 110,000 ha or 14% of
the area. These areas have high carrying
capacity for cattle, sheep, and alpaca. In
the areas near local markets, dual purpose
or dairy production is recommended. In
the areas with bofedales, alpaca
production is a better alternative. Sheep
constitute a flexible buffer alternative that
can be accommodated throughout this
AEZ. Current biomass production of less
than 5 t/ha and low quality might be
increased to more than 8 t/ha of good
qua! ity pasture (alfalfa, ryegrass, and
white clover) . lncrements on the order of
40% to 50% in gross income are feasible.
The third zone was classified as extensive
livestock production. With 51% of the
area, equivalent to 394,000 ha, this zone
367
......
..~
390000
20000
~0000
IS
Titicaca Lake
~..
1
..
!!
+
AQro ecologiQI zone
~ C1opl1hd Pastu1t
B
~..
.~
A
+
......
ICiti<d.cs
10
......
20000
SOOOO
20
368
Conclusions
The paper described new tools and methods to be incorporated into the AEZ
method. Through the inclusion of the
methods presented, scientists and decision
makers have access to a dynamic tool
for AEZ, even in data-scarce environments. The inclusion of remote sensing in
different parts of the method, together
with process-based climate interpolation models, add robustness to existing
procedu res.
In a practica! sense, several alternatives
have been assessed to improve the management of the natural resources of the
watershed. Since the work was jointly
executed with local professionals, the
chances to positively impact the sustainable management of the watershed are
greatly enhanced.
Acknowledgement
The Peruvian National Natural Resources
lnstitute (INRENA) kindly provided the soil
maps for the study area.
References
Aguilar, C. and R. Caas. 1991.
Produccin ovina para el Altiplano de
Puno, Per. Ciencia e investigacin
agraria 18:1 &2. p. 23-46.
Anderson, J.T., E.T. Hardy, J.T. Roach, and
R.E. Witmer. 1976. A land use and land
cover classification system for use with
remote sensor data: U .S Geological
225.
Arguelles, L. and R.O. Estrada (eds.). 1990.
Perspectivas de la Investigacin
Pecuaria para el Altiplano. Centro
Internacional de Investigaciones para el
Desarrollo / Proyecto de lnvestigacion
de Sistemas Agropecuarios Andinos.
Lima, Per. 504 p.
Baigorria, G.A., W.T. Bowen, and J.J.
Stoorvogel. 2000a. Climate/Weather
interpolation: A process-based spatial
interpolation model. Proceedings of
Annual meetings of American Society
of Agronomy - Crop Science Society of
America - Soi 1 Science Society of
America (SSA) held 5-9 Nov. 2000. SSA,
Minneapolis, Minnesota. p. 421.
(Abstract.)
Baigorria, G.A., J.J. Stoorvogel, and W.T.
Bowen. 2000b. Spatial-interpolation
rainfall model based on topography and
wind circulation. Proceedings of Annual
meetings of American Society of
Agronomy - Crop Science Society of
America - Soi 1 Science Society of
America (SSA) held 5-9 Nov. 2000. SSA,
Minneapolis, Minnesota. p. 421.
(Abstract.)
Bowen, W., H. Cabrera, V. Barrera, and G.
Baigorria. 1999. Simulating the response
of potato to applied nitrogen. In: lmpact
on a changing world. Program report
1997-98. lnternational Potato Center,
Lima, Peru. 458 p.
FAO (Food and Agricultura! Organization).
1997. Zonificacin agro-ecolgica,
Gua general. Servicio de Recursos,
Manejo y Conservacin de suelos.
Direccin de Fomento de Tierras y
Aguas, FAO. FAO, Rome, ltaly. 82 p.
Genin, D., H.-J. Picht, R. Lizarazu, and T.
Rodrguez. 1995. Waira Pampa: Un
sistema pastoril camlido-ovinos del
Altiplano rido Boliviano. IRD (lnstitut
369
370
J. StoorvogeP
Models of crop and soil systems are useful tools for understanding the complexity of the soil-plant-atmosphere continuum. Their application, however, is
limited by the availability of weather data that drives the processes described
in such models. In this study, we describe a process-based interpolation model
being developed for estimating maximum and minimum temperatures, precipitation, and solar radiation in mountain environments. The model is
parameterized with data obtained from three weather stations set along an
altitudinal gradient of 3020 to 3590 m above sea level in the La Encaada
watershed near Cajamarca, Peru. Using an independent data set from a
fourth station within the same watershed, we show that model estimates for
daily maximum and minimum temperatures agreed well with observed data.
The accuracy of model estimates for precipitation and solar radiation is still
being evaluated.
Plant growth and soil-related processes are
strongly influenced by weather. To simulate these processes accurately, models of
crop and soil systems require weather
data, which typically include daily values
for maximum and minimum temperatures,
rainfall, and solar radiation. Each of
these variables affects, to sorne degree,
processes such as photosynthesis, evapotranspiration, and the rate of plant growth
and development, as wel 1 as soi 1-related
processes that determine water and
nutrient availability.
One of the principal limitations to these
models is the lack of weather data. This is
especially true in tropical mountain
environments where few stations are set up
to record weather data. To overcome this
limitation, and to move towards the useful
application of models in the analysis of
1
371
372
Results
A predominant characteristic of climate in
the La Encaada watershed is illustrated
by the monthly means shown in Table 1.
That is, temperature falls and rainfall
increases with increasing elevation. For
maximum temperatures, the annual
average decreases from 1 6C at Manzanas
(3020 m) to 14C at Usnio (3260 m) to
11 C at La Toma (3590 m). Absolute trends
are smaller for minimum temperatures: the
annual average decreases from 6C at both
Manzanas and Usnio to 3C at La Toma. In
Figure 1, a comparison of daily temperature extremes recorded during 1999 also
shows more pronounced differences for
maximum temperature among sites.
Although the annual distribution of rainfall
varies somewhat, in most months, rainfall
tends to increase with elevation (Table 1).
The relationship with elevation is better
reflected in the average annual rainfall,
which shows Manzanas receiving a total
Table 1. Monthly means far daily maximum and minimum temperatures (TMAX, TMIN), rainfall (RAIN), and
global solar radiation (SRAD) recorded at stations in La Encaada placed along an altitudinal gradient
of 3020 m (Manzanas), 3260 m (Usnio) and 3590 m (La Toma) above sea level.1
Aug
Sep
Mar
Apr
May
Jun
Jul
Oct
Nov
Dec
Variable
Jan
Feb
TMAX (C)
11
12
12
12
11
12
11
10
11
La Toma
10
10
10
14
13
14
15
14
14
15
15
14
15
14
14
Usnio
17
17
15
15
16
16
18
18
16
16
Manzanas
17
16
TMIN (C)
2
4
3
3
3
3
4
3
3
2
3
La Toma
3
5
7
6
5
6
6
7
6
6
6
6
Usnio
6
4
5
7
6
8
6
Manzanas
7
7
7
7
5
4
RAIN (mm)
13
20
57
52
73 133
74
46
56
92 139
99
La Toma
11
75
21
9
40
65
72
70
43
Usnio
92 107 11 o
20
41
50
69
43
17
3
Manzanas
73
36
55 140 105
SRAD (MJ m2/d)
20
19
22
20
20
19
18
16
La Toma
19
15
18
18
22
22
18
15
18
19
19
16
Usnio
20
16
18
17
Manzanas
1 Manzanas
21
17
19
17
21
16
17
19
16
16
(7.118S, 78.310W); Usnio (7.089S, 78.316W ); La Toma (7.062S, 78.282W).
22
20
373
22
Manzanas
3020m
Usnio
3260m
La Toma
3590m
~
(])
:s
(ti
18
(;
c..
E
:::::
E
14
x:
ctS
~
10
50
100
150
200
250
300
350
Day of year
18
Manzanas
14
3020m
Usnio
3260m
La Toma
3590m
~
(])
:s
10
~
(])
c..
E
:::::
E
x:
ctS
:.?!
50
100
150
200
250
300
350
Day of year
Figure 1. Daily maximum and minimum temperatures recorded during 1999 at three stations located along an
altitudinal gradient in La Encaada, Peru.
37 4
18
~---------------.
- ..
TMAX=0.0093x+44.41
R2 = 1
........
....
.....
-- ..
- - .. -
3100
3200
3300
3400
3500
..
3600
Elevation (m)
375
10
9
y= 1.1256x - 0.8894
R2 =0.7419
8
7
~
"C
:5
E
;
1-
10
Figure 3. Relationship between minimum temperature (TMIN) observed at the Calvario station (3250 m) and that
estimated using only the temperature lapse rate (top figure) ora simulation model that incorporates the effect of
topographic parameters obtained from a digital elevation model (bottom figure). The broken line is the regression
line and the salid line is the 1:1 line.
376
Discussion
A preliminary analysis of the interpolation
model being developed for La Encaada
indicates that it can be successfully
applied to estimating the spatial variability of daily maximum and minimum
temperatures within the watershed. Further
analysis should focus on evaluating daily
estimates for rainfall and solar radiation.
The interpolation model described here
has recently been expanded to include
spatial estimates of rainfall based on
topography and wind circulation (Baigorria
et al., 2000).
A critica! question for future work is how
accurate such an interpolation model
might be for other watersheds where only
one or two weather stations exist, or where
a few weather stations are spread across a
larger region. In an attempt to address
these questions, we have formed a partnership with the Peruvian lnstitute for
Agrometeorology and Hydrology
(SENAMHI) that includes an evaluation of
the interpolation model using weatherstation data from throughout Cajamarca
Province.
Because few watersheds in the Andes have
even one weather station, another approach we are investigating is the use of
remote-sensing techniques to estimate
References
Baigorria, G.A. and W.T. Bowen. 2001. A
process-based model for spatial interpolation of extreme temperatures and solar
radiation. In: Proceedings of the Third
lnternational Symposium on Systems
Approaches for Agricultura! Development [CD-ROM computer file]. CIP,
Lima, Peru.
Baigorria, G.A., J.J. Stoorvogel, and W.T.
Bowen. 2000. Spatial-interpolation
rainfall model based on topography and
wind circulation. In: 2000 agronomy
abstracts. ASA/CSSA/SSSA, Madison,
WI, USA. p. 421.
De la Cruz, J., P. Zorogasta, and R.J.
Hijmans. 1999. A digital atlas of natural
resources in Cajamarca. Production
Systems and Natural Resources Management Department Working Paper
No. 2. CIP, Lima, Peru. 49 p.
de Scally, F.A. 1997. Deriving lapse rates
of slope air temperature for meltwater
runoff modeling in subtropical mountains: An example from the Punjab
Himalaya, Pakistan. Mountain Research
and Development 17:353-362.
Diak, G.R., W.L. Bland, J.R. Mecikalski,
and M.C. Anderson. 2000. Satellitebased estimates of longwave radiation
for agricultura! applications. Agricultura! and Forest Meteorology
103:349-355.
Glassy, J.M. and S.W. Running. 1994.
Validating diurnal climatology logic of
CIP Program Report 1999 - 2000
377
378
Eight viruses infect ulluco (Vllucus tuberosus Caldus), an Andean tuber crop.
Three of them, potato leafroll virus (PLRV), Andean potato latent virus, and
potato virus T, also infect potato (Solanum tuberosum L.). Arracacha virus A
and papaya mosaic virus, ulluco isolate (PapMV-U) infect other crops. PLRV,
one of the most damaging potato viruses, was detected in healthy potato
plants growing in the field next to PLRV-infected ulluco plants, indicating that
viruses can be disseminated among different Andean tuber crops under
natural conditions. The viruses with the highest incidence were PapMV-U and
three others, ullucus virus C, ullucus mild mottle virus, and ullucus mosaic
virus. In two field experiments in Junn, Peru, ulluco plants of accession MH290 infected with ullucus virus C or ullucus mosaic virus, and ulluco plants of
native variety Jaspeado infected with PLRV or PapMV-U yielded about 30%
less than the healthy control plants. Also, during field exposures, only a low
percentage of healthy plants were reinfected with viruses. These results
indicate that the production and use of virus-free ulluco seed tubers is justified in this environment and, therefore, the use of better seed by farmers has
been promoted, particularly in the La Libertad community in Junn, Huancayo
Department.
381
382
Table 1. lncidence of ulluco viruses in accessions from different countries and from departments of Peru
maintained at CIP's in vitro germplasm collection and in plants from farmers, fields (Junn and
Huancavelica, Peru).
Samples
from:
In vitro
collection
Field 2
lnfected {%}1
Accessions
uve UMV PapMV-U UMMV PLRV APLV
{No.}
Argentina
39
76
45
90
66
21
59
Bolivia
77
66
46
62
22
52
22
4
Colombia
90
60
90
90
30
90
Ecuador
5
39
26
50
90
o 26
Peru
258
74
61
65
39
29
35
Apurimac
4
30
o 60
o 30
90
o
Amazonas
3
o
o
o
35
35
Ancash
25
90
66
39
31
27
66
Ayacucho
8
45
o
o 21
60
38
68
33
16
30
Cajamarca
63
76
61
o o
o
o 45
o
Cerro de Paseo
2
82
93
65
73
36
41
43
Cusca
Huanuco
2
o 90
o
o
o
90
o o
o
Junn
1
o
o
o
o
La Libertad
6
65
45
o 35
45
o 22 32
7
68
57
32
Lima
6
90
o 35
90
90
Piura
90
74
Puno
38
77
74
61
25
33
Total
383
72
65
40
31
{%}
57
37
Country
Department
Peru
Ju nin
Chicche
S.Juan deJarpa
Huaracayo
Huancavelica
Pazos
Total
{%}
PVT
9
16
AVA
15
14
o
o
o o
o o
o o
o o
o 12
o
21
o o
o o
o
13
o o
o o
o o
o o
o o
o
33
42
43
41
68
62
56
65
62
51
49
51
o
o
21
24
17
nt 3
nt
nt
o
o
o
72
57
43
58
24
nt
67
52
42
61
22
180
180
180
72
180
720
383
Oca, mashua
Potato
Po tato
Potato, oca, mashua
Arracacha
Germplasm collection from UNMSM (Universidad Nacional Mayor de San Marcos, Lima, Peru) and CIP (lnternational
Patato Center, Lima, Peru).
2 Ali virus isolates, except PLR\/, were mechanically transmitted.
3 Virus transmitted by true seed repo rted in other hosts, but not yet confirmed in ulluco.
4 PLRV was isolated from the naturally infected accession and maintained in the same plant.
Results
Virus identification and viral incidence
Ei ght viruses infect ullu co (Table 2). Of
th ese, viruses PLRV, APLV, PVT, and AVA
384
Table 3. Reinfection (%) 1 of virus-free ulluco Jaspeado plants in farmers' fields in La Libertad, Junn, Peru
(3500 m).
Field
Planting season
1996-1997
Farmer 1
Farmer 2
Farmer 3
PLRV
APLV
UMV
uve
1.3
1.3
1.3
1.3
2.5
2.5
2.5
2.5
PLRV
10.0 6.6
PLRV
2.5 3.5
1997-1998
1998-19992
Not planted
Not planted
UMV
uve
PapMV-U
UMMV
PLRV
APLV
Not
5
2.5
1.3
2.5
11
2.5
4.8
3.5
2.5
3.5
7.0
3.5
UMV
uve
PapMV-U
UMMV
PLRV
APLV
~lanted
27.5 9.8
12.5 7.2
5.0 4.8
13.8 7.6
51 .3 11 .0
13.8 7.6
Not
~lanted
Table 4. Reinfection (%) 1 of Jaspeado ulluco plants and seed tubers from virus-free ulluco produced in field
in La Libertad, Junn , Peru (3500 m).
Samples 2 Categor'f Evaluated
UVC
UMV PapMV-U
UMMV
PLRV
APLV PVT AVA
Plants
Virus-free
48
O
O
O
O
o
o nt 4 O
Basic
105
43.7
43.7
O 43.7 41 9.5 11 6.0 nt
O
eertified
58
128.5 128.5
34.5
97.5 4813.1 2811 .8 nt
O
Tubers
eertified
49
3313.4 4514.2
O 2712.7
o 5714.1 o o
Note: See Table 2 far full names of viruses.
1 confidence limits (p = 0.05).
2
Plants and tubers from cropping season 1997-1998 and 1999-2000, respectively.
Viru s-free = 1st generation in screenhouse; Basic = 2nd generation in field; Certified
4 nt = not tested .
385
13.9 kg
Discussion
O ur obse rv ati ons and those from To ledo et
al. (199 4) indi ca te th at v iruses u ve,
PapMV-U, UMV, and UMMV are
w idespread in ulluco (Tab le 1). Hi gh viral
in cidence has also been repo rted in
Bo li v ia and Ecuado r (D uqu e and H ermann ,
1994; Badani et al., 1997). Ei ght viru ses
have bee n fo und in fec tin g ullu co . Viral
incidence data of PLRV, APLV, PVT, and
AVA were not ava il ab le befare this stud y.
Th e info rm atio n co mpil ed here on th e
identificat ion and distributi on of viru ses in
ullu co permits recog nizin g dissemin atin g
viru ses and those restri cted to ce rtain
geographi ca l areas (Tab le 1). lt is of
ep id emio log ica l interest that PLRV, A PLV,
PVT, and AVA naturall y infect other
Andean crops, espec iall y patato (Tab le 2)
(Janes and Kenten, 1978; Li z rraga et al. ,
1997; Li z rraga et al. , 2000). Farm ers'
trad itio nal croppin g systems (ulluco and
potato in mi xed crop ping, ullu co fields
beside patato fields, or ulluco pl anted in
fields w here patato was prev iou sly grown )
favor vi rus dissemination betwee n crops.
Th at could be occurring with PLRV, one of
the most damag ing patato viru ses, and thi s
study suggests th at, in Peru, potato seedproducing fi eld s should be far from both
patato and ullu co f ield s. Under expe rim ental co nditi ons, PLRV was transmitted by
th e gree n peach ap hid (Myzus persicae)
19.8 kg
14.1 kg
Figure 2. Effect of viruses on yield of ulluco Jaspeado in secondary infection with PLRV and PapMV-U. Tubers
harvested from 20 plants per each treatment.
386
Table 5. Average yield of tubers (kg) from ulluco plants (MH-290 and Jaspeado) with secondary infection
(seed tubers were virus infected), planted in two different cropping seasons and places in Junn, Peru.
Cropping season,
Treatment
Yield 1
Yield
place and variety
Total weight
Commercial quality
reduction (%) 2
1995-1996
Healthy control
26.9 a
16.0 a
(Huancayo)
UMV
19.1 be
10.5 be
29
27
MH-290
uve
19.7 be
10.7 be
PapMV-U
24.2 a
14.5 ab
10
UMV +uve+ PapMV-U
16.7 be
10.5 be
38
Healthy control
19.8 a
nt 3
1998-1999
UMV
16.4 a
nt
17
(La Libertad)
uve
19.5 a
nt
2
Jaspeado
PapMV-U
14.1 b
nt
29
PLRV
13.9 b
nt
30
Note: See Table 2 for full names of viruses.
1 Means within columns followed by the same letter do not differ significantly (p = 0.05). lnner rows of 66 and 20
plants for MH-290 and Jaspeado, respectively.
2 Compared with total yield of healthy control.
3 nt = not tested.
Table 6. Average yield of tubers (kg) from ulluco Jaspeado, planted in La Libertad, Junn, (cropping
seas ons 1997-98 and 1998-99).
Yield 1
Cropping
Treatment
Yield
reduction (%)2
sea son
Total weight
Commercial quality
1997-98
Basic
18.90 a
13.19 a
lnfected 3
14.11 ab
10.27 ab
22
From farmer
13.35 b
9.39 b
29
Positive selection
9.86 b
7.86 b
40
1998-99
Virus-free (5 g)
37.27 a
28.96 a
Virus-free (25 g)
36.79 a
25.09 ab
13
eertified (5 g)
35.66 a
19.62 e
32
eertified (25 g)
34.09 a
20.95 be
28
lnfected (5 g)
36.80 a
19.52 e
33
lnfected (25 g)
31.98 a
18.74 e
35
1 Means within columns followed by the same letter do not differ significantly (p = 0.05). Plot of 40 plants.
2 Compared with commercial quality yield of healthy control.
3 lnfected with at least one of following viruses: UMV, UVC, PapMV-U, and UMMV.
387
Conclusions
388
Acknowledgments
This work is part of the collaborative
program Biodiversity of Andean Root and
Tuber erops funded by the Swiss Agency
for Technical eooperation. Thanks to
R. Burns of the Scottish Agricultura!
Science Agency, Edinburgh, Scotland, UK,
for supplying PVT antibodies, and R.
Woods from Rothamsted Ex peri mental
Station, Harpenden, U K, for AVA antiseru m. Sorne of the tables, with additional
information, have been translated from
Spanish from Lizrraga et al. (1999) with
the kind permission of Fitopatologa.
References
Badani, A.C., S. Conzales, C. Plata, and
E.N. Fernandez-Northcote. 1997.
Incidencia de virus en papalisa (Ul/ucus
tuberosus Loz) en Cochabamba, Bolivia.
In: IX Congreso Internacional de
Cultivos Andinos, Libro de resmenes.
Universidad Nacional de San Antonio
de Abad del Cusco (UNSAAC), Centro
de Investigaciones en Cultivos Andinos
(CICA), and Asociacin ARARIWA,
Cusco, Per. p. 15.
Bertschinger, L. 1992. Modeling of potato
virus pathosystems by means of quantitative epidemiology: An exemplary
case based on virus degeneration
studies in Peru. Ph.D. thesis. Swiss
Federal lnstitute of Technology, Zurich,
Switzerland. 111 p.
Brunt A.A., S. Phillips, R.A.C. Jones, and
R.H. Kenten. 1982. Virus detected in
Ullucus tuberosus (Basel laceae) from
Peru and Bolivia. Annals of Applied
Biology 101 :65-71.
Duque, L.M. and M. Hermann. 1994.
Erradicacin de virus en melloco
(Ul/ucus tuberosus Caldas). Carta de
SEFIT - Sociedad Ecuatoriana de
Fitopatologa 3(6):1-3.
Hodge, W.H. 1951. Three native tuber
plants of the high Andes. Economic
Botany 5:185-201.
Jones, R.A.C. and R.H. Kenten. 1978.
Arracacha virus A, a newly recognized
virus infecting arracacha (Arracacia
xanthorrhiza: Umbelliferae) in the
Peruvian Andes. Annals of Applied
Biology 90:85-91.
Lizrraga, C., M. Querci, M. Santa Cruz,
l. Bartolina, and L.F. Salazar. 2000.
Other natural hosts of potato virus T.
Plant Disease 84:736-738.
Lizrraga, C., M. Santa Cruz, J.L. Marca,
and L.F. Salazar. 1999. La importancia
de los virus que infectan a Ullucus
tuberosus Caldas en el Per.
Fitopatologa 34:22-28.
Lizrraga, C., M. Santa Cruz, and
L.F. Salazar. 1997. Progress in identify-
389
Oca is an edible starchy tuber grown in the Andes. To improve its culinary
quality, harvested oca is typically exposed to direct sunlight (sunned) for
several days prior to consumption. Five native cultivars were examined to
determine the changes in nutritional composition of oca as a result of sunning.
This practice increased the proportions of dry matter, soluble solids, and
sugars (mainly sucrose) and decreased total acids, due mainly to a large
reduction in malate and glutarate. High levels of oxalate, an anti-nutritional
factor, were found, ranging from 306 to 539 and 251 to 451 mg/100 gin
edible matter of freshly harvested and sunned tubers, respectively. Sunning
reduced oxalate levels in dry matter by an average of 26%. Further study will
be necessary to fully assess the food safety of oca and to provide recommendations for maximum daily intake.
391
Results
Although their edaphic space was restricted by being pot-grown, the oca plants
developed normally, showing a typical
ontogenetic sequence (slow initial development fol lowed by vigorous shoot and
foliage growth, flowering, tuber bulking,
and senescence). Plant and final tuber size
and plant habit were comparable to those
usually observed for oca under field
conditions.
During sunning, maximum tuber flesh
temperatures reached 35C in the early
Latitude Longitude
Comments
'C
Q)C")OQf'-
~~~~;;~~~~~
......
-.::j"'
C\J
"I";'"
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C\J C\J
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C.Nr--.:t-c.oC\JOOOC.OC\J~O~
~~~
LL
C.0
et')
C\J~
C.O C.0
C") et')
L{)
C")~~
C\J
a>c a.M. t -.
e: OMO>OC.OMC\JM-.::l"'OO>-.::l"'C.0
e: C\J
t - L{) C\J O> O> C\J e.o 00 e.o L{)
::::11
OOO>OOOC.OC\JC.OMM
(1)
L{) et') ~
"'O
cu
Cl
o
o
e::
o
E
en
C2.
E e:
o(.) ::J
.l!:S Cl Cl Cl Cl Cl Cl Cl Cl Cl Cl
ClCl~EEEEEEEEEE
(.)
.E
cu
..e::
(.)
e-.: cu
:e
-~
:e
ca ca
Q)
t-
>
393
Discussion
This study is the first to provide extensive
data on the carbohydrate and organic acid
composition of freshly harvested oca
tubers. lt is also the first to address the
compositional changes associated with
sunning, a traditional postharvest technique to render the tubers more palatable.
The oca cultivars used in this study are
morphologically diverse and represent
different agro-ecologies found in Argentina, Ecuador, and Peru. These data are
therefore not constrained by using geographically limited material and allow
inference to be made concerning oca
throughout the species range.
In agreement with earlier studies (Cortes,
1978; Cross et al., 1989; Kays et al.,
1979), oca was shown to consist mostly of
HN-1146
34
106
116
88
102
37
24
27
54
85
89
References
Cortes H. 1978. Avances de la
investigacin en oca. In: Tapia, M.E.
and M. Villarroel (eds.). Proceedings of
the First lnternational Congress on
Andean Cultivars held 25-28 October
1977 in Ayacucho, Peru. lnter-American
lnstitute for Cooperation on Agriculture,
La Paz, Bolivia. p. 227-243. (in
Spanish)
Cross, R., F. Koch, l. Malaga, A.F. de
Miranda, H. Schoeneberger, and
CIP Program Report 1999 - 2000
395
396
Downy mildew (Peronospora farinosa (Fr.) Fr) is the most important disease of
quinoa (Chenopodium quinoa Willd.) in the Andes. To quantify the impact of
the disease on grain yield under natural conditions, a field experiment with
eight quinoa cultivars with and without application of fungicides, was carried
out in Huancayo, Junn, Peru (3200 m), where the crop is grown traditionally.
Area under the disease progress curve (AUDPC) values were calculated,
based on evaluations of disease severity (percentage leaf area affected). The
cultivar Utusaya, originating from the Bolivian salt desert (200 mm annual
rainfall), was strongly affected by downy mildew, which caused complete
defoliation, premature maturation, and a yield loss of 99%. Even in the most
resistant cultivar, yield was reduced 33%, indicating the destructiveness of
this disease.
397
Experimental design
The tria! was carried out in the experimental field of CIP, Huancayo, Junn, Peru,
(3200 m) during the 1999/2000 growing
season under conditions of natural downy
mildew infection.
Cultivars
Eight quinoa cultivars with different
geographical origins were included in the
study: Utusaya (salt desert); LP-48, La
Molina 89 (coast); Blanca de Juli,
Kancolla, and Jujuy (highlands); and
Amarilla de Marangan and lngapirca
(val ley). Utusaya and LP-48 are the
earliest maturing cultivars; La Molina 89,
Amarilla de Marangan, and lngapirca the
la test.
AUDPC
= ~, ( Y;+
398
Harvest
Seed yield was estimated from harvest of
the four center rows. The number of plants
in each plot was counted. Harvest was
done when the plants were mature,
starting 131 DAS with Utusaya and LP-4B,
and ending 165 DAS with La Malina 89,
Amarilla de Marangan, and lngapirca.
Amarilla de Marangan
100 - . - - - - - - - - - - - - - - - - - - - .
?f.
- - Untreated
__._ T reated
80
Cll
!!!
Cll
1ii
60
"O
40
:g
20
.!l1
69
Statistical analysis
The effect of cultivar and fungicide on
yield and AU DPC, and the effect of
fungicide treatment on the number of
plants per plot were analyzed by ANOVA.
The correlation between yield and AU DPC
was analyzed with correlation analysis
(SAS System version 6.12, SAS lnstitute
lnc., Cary, NC).
98
131
?f. 80
Cll
!!!
Cll
60
1ii
.!l1
"O
40
20
Results
98
69
131
DTreated
Untreated
400)
oa.. :mo
o
::>
2CXXl
<(
1CXXJ
<tl
en
::::>
=>
::::>
--,
Q)
"O
ro
>-
""'!"
::-
o....
_J
::::>
--,
oo
e
<tl
::,,::::
-~o.
<tl
Ol
<tl
in
:2
Q)
"O
;::
C1l
00
<tl
<tl
:2
_J
<(
<tl
Cultivar
399
Table 1. Seed yield and AUDPC values in eight quinoa cultivars with two levels of downy mildew (with and
without fungicide treatment).
AUDPC 2
Yield (kg/ha) 1
Cultivar
3
3
Untreated Yield loss (%)
Treated
Untreated
Difference
Treated
Salt desert cultivar
579
2106
23
4636
4057 ***
Utusaya
99 ***
Coastal cultivars
4092
1021
384
2790
LP-48
75 ***
2406 ***
7686
3263
256
852
La Malina 89
58 ***
596 *
Highland cultivars
1465
Blanca de Juli
5891
510
2976
75 ***
2466 ***
3083
408
1829
Kancolla
4748
35 *
1421 ***
1846
544
2478
J~uy
5424
66 ***
1934 ***
High altitude valley cultivars
Amarilla de M.
7559
5073
33 ***
303
882
579 *
144
lngapirca
6570
3562
46 ***
1207
1063 ***
Leve! of significance: * P = 0.05, *** P = 0.001 (ANOVA)
1 The harvested area of each plot was 8.4 m2. The yield was converted into kg/ha by multiplying g/parcel with a factor
1.19.
2 Area under the disease progress curve.
3 The plots were sprayed with metalaxyl (systemic) and mancozeb (contact) to control downy mildew.
Acknowledgments
We are grateful to Carlos Hualhuas and
Lorenzo Safra for handling the seeds.
Thanks to Felipe Mendiburu for helping
with the statistics. This study was supported by the Danish government (RUF
grant no. 90929).
References
Alandia, S., V. Otaz, and B. Salas. 1979.
Enfermedades. In: Tapia, M.,
H. Gandarillas, S. Alandia, A. Cardozo,
A. Mujica, R. Ortiz, V. Otaz, J. Rea,
B. Salas, and E. Sanabria, (eds.).
Quinua y Kaiwa. Editorial llCA,
Bogot, Colombia, p. 137-148.
401
403
404
156
o
397
13,582
2134
Potassium (mg/I)
Magnesium (mg/I)
Sodium (mg/I)
Nitrates(mg/1)
pH
EC (mS/cm)
469
1120
7018
37
7
43
405
(A)
(A)
~
1
~
~
(BC)
(BCD)
(BCD)
(CDE)
(CDE)
~
1
'1
,,
,,,,,,,,,
:-...-...................................
.............
........................................................................
(DE)
~
1
Amaranth
Osear Blanco
(E)
SI
(E)
(F)
Quinua
03-26-0036
1
~
'1
~'''''''"
1
1
0.1
0.2 0.3
0.4
0.5 0.6
0.7
0.8
Quinua
Utusaya
1
0.9
1.1
Figure 1. Salinity tolerance according to TS/TC values in quinoa varieties (03-26-0036 and Utusaya) and in the
amaranth variety Osear Blanco.* = Measured at anthesis. A-F indicate significant differences at the 95% level as
an average between the two quinoa varieties and the one of amaranth. (dm = dry matter; MPa = MegaPascal)
(seconds per centimeter) under salinity of
7.77 mS/cm and from 12.2 to 28.2 s/cm at
11.88 mS/cm; increasing salinity tended to
increase this resistance.
Conclusion
Amaranth demonstrated very 1ittle abi 1ity
to regulate leaf water potential and
stomatal conductivity, and the plants died
at high salinity levels. According to the
TS/TC factor, amaranth was four times
more susceptible to salinity than quinoa in
relation to seed yield. Quinoa demonstrated the ability to accumulate salt ions
in its tissue to control and adjust leaf water
potential. That enabled the plants to
maintain cell turgor and limit transpiration
under saline conditions, thus avoiding
406
References
Bosque, H.D. 1998. Ecophysiological
analysis of drought and salinity stress of
quinoa (Chenopodium quinoa Willd.).
MSc thesis in Soi 1 Science and
Eremology. Faculty of Science, Faculty
of Agricultura! and Applied Biological
Sciences, University of Gent.
lnternational Center for Eremology,
Gent, Belgium. 122 p.
Brady, N.C. and R.R. Weil. 1999. The
nature and properties of soils. 121h
edition. Prentice-Hal 1, lnc, Upper
Saddle River, NJ, USA.
Canahua, A. and Z. Cutipa. 2001.
Produccin de la quinua en waru-waru:
Perspectivas y limitaciones. In:
Jacobsen, S.-E. and Z. Portillo (eds.).
Memorias, Primer Taller Internacional
sobre Quinua - Recursos Geneticos y
Sistemas de Produccin, 10-14 May
1999, Universidad Nacional Agraria La
Molina, Lima, Peru. CD-Rom available
from CIP, Lima, Peru.
Cari, A. 1978. Efectos de la salinidad y
fertilizacin potsica en dos variedades
de quinua (Chenopodium quinoa Willd).
MSc thesis. Universidad Nacional
Tcnica del Altiplano, Puno, Peru. 66 p.
Flowers, T.J., M.A. Hajibagheri, and
N.J.W. Clipson. 1986. Halophytes. The
Quarterly Review of Biology 61 (3):
313-337.
Flowers, T.J. and A.R. Veo. 1986. Ion
relations of plants under drought and
salinity. Australian Journal of Scientific
Research 13:75-91.
Jacobsen, S.-E. and A. Mujica. 2001.
Avances en el conocimiento de
resistencia a factores abiticos adversos
en la quinua (Chenopodium quinoa
Willd.). In: Jacobsen, S.-E. and
Portillo (eds.). Memorias, Primer
Taller Internacional sobre Quinua Recursos Geneticos y Sistemas de
Produccin, 10-14 May 1999,
Universidad Nacional Agraria La
Molina, Lima, Peru. CD-Rom available
from CIP, Lima, Peru.
z.
407
408
411
413
414
Developing interventions
After the data collection phase, algorithms
were devised following the flow diagram
in Figure 2 to divide the watershed into
soil conservation zones. In collaboration
with the PRONAMACHCS team and
published tables (PRONAMACHCS, 1998),
a set of rules was established to identify
potential soil conservation nterventions
(no intervention, sloping terraces, infiltration ditches, and reforestation) on cropped
land (Table 1).
There was a consensus that soi 1 conservation measures were not of great
importance for slopes of less than 5% and
that no interventions other than vegetation
restoration and forest plantations were
possible on slopes of more than 40%. The
main interventions would focus on the
intermediate slope types, which were
categorized into two groups: 5-15%
(terracing possible) and 16-40% (infiltration ditches, terracing only possible if the
soils are deep).
The six ground cover classes originally
used in the data collection phase were
simplified to three, based on 'permanency'
of ground cover: zones of permanent
vegetation (grasslands, forest), zones of
annual disturbance (annual crops plus
associated short-term fal low), and degraded zones where little vegetative cover
remained. With respect to crop type,
cereal (maize, wheat, barley, oats), tuber
(potato, oca, ollucu), and legume (peas,
76'
68'
72'
o
Colombia
Ecuador
4'
4'
CAJAMARCA
Department
Brasil
Altitude [m]
CJ 1-1000
CJ 1001 - 2000
c---12001 - 3000
CJ 3001 - 4000
-
4001-5000
CJ >5000
12'
12
.~
.;:::
oco
16'
16'
so
72'
76'
68'
415
Elevation
model
Slope
Slope
classes
Ground
cover
Ground
cover
classes
Soil
1-------11---
Eros ion
risk
1----...__~soil depth1-------1~~----'
classes
Basemaps
C=>
Decision table
Results
Process
Proposal at
commnunity
level
Table 1. Decision rules far soil conservation interventions in annual cropping areas.
Soil depth
Erosion potential
Proposed intervention
Shallow<60 cm
Low
No intervention
Medium
Sloping terraces
High
lnfiltration ditches
Very high
Re-establish native vegetation
Deep>60 cm
Low
No intervention
Medium
Sloping terraces
High
Sloping terraces/infiltration ditches
Very high
Commercial forests
416
417
795000
805000
800000
810000
Land use
Annual
aopp;ng
Degraded
1D
Zonification
kea (ha)
No intervention
444
lnftration ditches
1655
Sloping terraces
2422
186
Commercial forest
Nativeforest
446
Rehabilitation
1085
9503
9215000
9215000
9210000
9220000
9210000
i!!!!!!'!"'!1iiiii;;;;ij2~!!!!'!"
aooooo
795000
805000
810000
776000
780000
Area ha)
124
1241
483
70
1662
9192000
9192000
75
4467
(\) Community boundaries
9188000
9188000
9184000
9184000
kilomeler.;
1
Praeclil:xlllTMi:one 17s
772000
776000
780000
Conclusions
Digitizing existing maps and
georeferencing sorne additional field
measurements can result in useful databases for natural resource management.
What is equally important is that more and
more local organizations (government
agencies, NGOs, university laboratories)
are acquiring the software and experience
necessary to apply and improve the
methodology. At the level of a watershed
like La Encaada or Asuncin, the methodology allows municipal and agency
personal to identify soil conservation
intervention zones or zones suitable for
new production systems. At the local
community level, the system is being
adapted to help envision and salve local
problems, especial ly around water issues.
References
De la Cruz, J., P. Zorogasta, and
R. J. Hijmans. 1999. Atlas digital de los
recursos naturales de Cajamarca.
Department of Natural Resource
Management working document No. 2.
lnternational Patato Center, Lima, Peru.
49 p.
Delgado Loayza, R. 1998. Evaluacin del
recurso hdrico de la microcuenca de
Ro Asuncin-Cuenca del Jequetepeque.
ASPADERUC/CONDESAN. Cajamarca,
Peru. 43 p.
Farrington, J. and C. Lobo. 1997. Scalingup participatory watershed development
in India: Lessons from the lndo-German
Watershed Development Programme.
Natural Resource Perspectives 1 7:1-6.
Jimnez Medina, M. 1996. Estudio
de suelos de la Microcuenca "La
Encaada" (semi-detallado).
ASPADERUC/CONDESAN, Cajamarca,
Peru. 73 p.
Jimnez Medina, M. 1998. Estudio de
suelos-Distrito de la Asuncin (semi-
419
420
1975-1996
C.B. Bussink and R.J. Hijmans
We used land-use maps for different years and areas in the Cajamarca catchment in the northern Peruvian Andes to study changes in land use between
1975 and 1996. Despite population growth, agriculture has not been intensified nor extensified. Neither did we find evidence for crop encroachment into
the higher parts of the catchment. The agricultural area has decreased and
the proportion of fallow land has increased. There has been a significant
increase of shrubs and bare soil at the expense of natural pastures, perhaps
indicating overgrazing. The area planted to specific crops has fluctuated, but
there has been a clear decrease in the area planted to barley and an increase
in wheat. The area with exotic tree species has also increased.
421
7'00'
Cajamarca
catchment
kilometers
5 10
o
!'8"1.5'
rJO'
78"{):}'
Figure 1. Location of the Cajamarca catchment in Peru and the study areas: 1975/78 (the whole catchment),
1991 (A), 1992/1993 (B), 1996 (C), and the zone with three studies (D).
Near Cajamarca town, at 2650 m, average
daily temperature is about 13C and yearly
precipitation is 720 mm. The temperature
is rather constant over the year, while
diurna! differences are about 17C. There
is a clear rainy season between October
and April (De la Cruz et al., 1999). The
altitude in the catchment is between 2000
m to 4200 m, with about 79% between
2600 m and 3800 m. The distribution of
crops by altitude has been used to distinguish main agroecological zones: the
cu ltivated pasture zone (irrigated val ley
bottoms), the maize zone (lower slopes),
the tuber zone (higher slopes), and the
natural pasture zone (jalea) (Kohler, 1986;
Kohler and Tillman, 1988; Seifert, 1990;
422
Results
Land use in 1975/78
423
Crops
Fallow
30
45
37
35
34
37
33
39
35
5
9
12
9
9
7
8
9
9
Bare soil
& rocks
9
10
15
9
11
10
12
9
12
Land use
Forest
Cultivated pasture
Maize
Wheat
13 Zone A (1975178-91)
Barley
mZone C (1975178-96)
C Zone B (1975178-92/93)
C Zone A+B+C
Rye& Oats
Pulses
Tubers
Fallow
Bare soil I rocks
-25
-20
-15
-10
-5
10
15
20
Figure 2. Change in land use far three zones with different time series (1975/78-1991, 1975/78-1992/93, and
1975/78-1996).
424
Table 2. Change in land use from one class to another between 1975/1978 and the 1990s (expressed in
percent of area of each land-use class in 1975/78).
1990s
Cultivated
Natural
Bare soil /
Shrubs
Forest
Crops
Fallow
Total
1915na
pasture
pasture
rocks
Natural
15
11
39
20
6
3
5
100
pasture
12
100
100
73
11
66
100
10
15
19
43
100
12
15
57
100
Shrubs
69
Forest
69
10
Cultivated
pasture
Crops
Fallow
Bare soil J
rocks
60
50
40
01975(78
30
1990s
20
10
425
Relative area
01975/78
1991
[!11996
Land use
Figure 4. Area of land use classes far three years (1975/78, 1991 and 1996) in one zone (Zone D; Figure 1).
farm size, only cover 10% of the cultivated area (Seifert, 1990). The decrease of
area planted to tubers and the increase of
area under cultivated pasture coincides
with Frias's (1995) study for the whole
Cajamarca Department. However, instead
of the decrease in wheat described by
Frias (1995), we found an increase.
The largest change is an increase in shrub
land and bare soil, mainly at the expense
of natural pasture. Overgrazing may have
led to soi 1 eros ion and loss of soi 1 ferti 1ity,
which in turn, may have led to an increase
in unpalatable shrubs. Alternatively, it
might be a sign of vegetation recuperating
because of a decrease in grazing pressure.
The i ncrease in forested areas match es the
pattern of agricultura! dis-intensification
and retraction. Also, over the past few
decades, there has been considerable
public investment in forestation in
Cajamarca, particu larly through a local
non-governmental organization.
One complication with our data is that we
compared not only across different years
but also across different zones. The only
zone for wh ich we had data for three
periods was zone D. However, it appears
that for sorne land-use classes, the effect
426
Acknowledgements
We thank Jorge de la Cruz, Felipe de
Mendiburu, Edwin Rojas, and Percy
Zorogasta for assistance; Jiefar Diaz,
Wilfredo Poma (UNC), and Pablo Sanchez
(ASPADERUC) for providing access to old
land-use maps; CONDESAN for partly
funding this research; and Joshua Posner
and Dominique Herv for reviewing of this
paper.
References
Chiln C., M.I. 1993. Estudio del uso
actual de la tierra de la intercuenca del
rio Sambar-Distrito Baos del Inca.
Master's thesis. Universidad Nacional
de Cajamarca, Peru.
De la Cruz, J., P. Zorogasta, and R.J.
Hijmans. 1999. Atlas digital de los
recursos naturales de Cajamarca.
Natural Resource Management Working
Document No. 2. lnternational Potato
Center, Lima, Peru. 49 p.
427
428
(2): 319-339.
GILB: An Update
The Global lnitiative on Late Blight (GILB) brings people together with the aim of
increasing and enhancing global effarts devoted to solving the problem of late
blight in patato, the world's most costly biotic constraint to patato production.
GILB was initiated in early 1996 when farty-one participants from developing and
industrialized countries met at a Project Design Meeting at CIP headquarters in
Lima, Peru, to plan a three-phase, ten-year program with specific priorities far
each phase. lnterest in GILB has grown rapidly, to the point that, in the year
2000, the GILB newsletter reached more than 633 late blight workers in over 77
countries.
GILB priorities were further modified and refined during a 1999 GILB-organized
global conference with 165 participants from 40 countries. Participants reviewed
the list of priorities established in 1996 and analyzed achievements under those
priorities. Because impressive progress had been made toward meeting the initial
priorities, in part through rapid advances in molecular technologies, new priority
areas were established to guide GILB activities far the next three years. The
highest-priority areas far GILB are breeding far host resistance, studying the
pathogen and host-pathogen relationship, promoting integrated pest management
(IPM), and providing training and infarmation.
Members at the GILB'99 Conference agreed that infarmation is a critica! need.
Therefare, GILB has concentrated its efforts on developing a Global Late Blight
lnfarmation System available online at the GILB web address (www.cipotato.org/
GILB). This online system includes
databases on Phytophthora infestans populations and their characterizations,
a directory of people interested and working in particular areas (indexed by
geographic area and theme),
a P. infestans bibliography,
links to other potato late-blight-related websites,
laboratory techniques,
avai lable research materials, and
links to relevant research databases.
A global catalog of late blight resistant patato varieties is being compiled and
will be available both online and as a CD-ROM. The catalog will include data
such as synonyms, parentage, breeder, releasing institution and date, whether it is
protected or not, source (where it can be obtained), and area of resistance (countries).
GILB promotes high-priority research by improving communications between and
among researchers and institutions and encouraging the transfer of technology.
Through GILB, research is facilitated by improving access to infarmation to late
blight working groups, especially in developing countries. The Global Late Blight
429
lnformation System and the GILB newsletter, along with sponsored symposia,
meetings, and workshops supported by GILB, all contribute to attaining these
goal s.
GILB members will next meet at the GILB Global Conference 'Late Blight:
Managing the Global Threat' to be held 11-13 July 2002, in conjunction with the
1 Sth Triennial Conference of the European Association of Potato Research (EAPR)
which will take place 14-19 July 2002 in Hamburg, Germany.
This report, particularly in the first seven papers in the potato section, show the
preadth and depth of CIP's research on Late Blight.
430
431
433
434
435
436
437
by
438
439
SIUPA
The CGIAR Strategic lnitiative on Urban
and Peri-urban Agriculture (SIUPA) has
selected Metro Manila as one of its global
network of field research sites. SIUPA has
worked closely with the CPT of the
440
llRDEP
Socio-economics and marketing
Food safety
Production systems
Environmental protection
Input management system
SIUPA
Supply of perishable/processed products
Urban agriculture and livelihoods
Environment and health
Agriculture and non-ag uses
Cross-cutting themes
Saving biodiversity
441
Table 4. Mapping IRRDEP projects using the SIUPA program framework (Campilan, 2001).
Theme
Household/community
lnstitutional
Policy
Supply of UPA
Development of year-round vegetable
products
production systems.
Development of fertilizer, growing media.
Edible landscaping.
Receptacle/container gardening.
lmprovement of varieties of vegetables.
UPA and
Development of marketing strategies
Development of marketing
livelihoods
and assessment of product flow.
strategies and assessment
of product flow.
Environment and Analysis far toxic heavy metals.
health
Assurance of toad safety in edible
crops.
Detection of pesticide residues.
Collection and selection of crop
varieties.
Recycling of river and sewage water.
Development of odorless compost
making.
Agricultural/nonagricultural uses
Cross-cutting
442
References
Asiaweek. 1999. Asiaweek best cities
1999. http://www.asiaweek.com/
asiaweek/features/asiacities/acl 999/
data/manila.html
Banal, Conrado R. 2001. Breaktime.
Philippine Daily lnquirer, 20 February
2001 issue. p. 82.
443
This study assesses past, present, and future development trends of the dairy
sector in the surroundings of Lima, Peru. Results show that peri-urban milk
producers currently enjoy favorable market conditions, thanks to increased
competition among local milk buyers, who seek to exploit spare processing
capacity. Yet, this situation is likely to change in the near future as milk
production rapidly expands. The resulting drop in price will be likely to put
many small and medium farms out of business, whereas large farms will be in
a better position to confront lower prices, as a result of considerably lower
average production costs (economies of scale) and higher productivity,
mainly due to better feed quality. The only viable strategy for small and
medium milk producers is herd growth and the creation of producer associations to strengthen their market position versus suppliers of inputs and
purchasers of milk. Nonetheless, improvements in feeding and reproduction
practices will be essential to achieve faster herd growth. For large farms, the
key to improved profitability will be to efficiently manage the different
contracted specialists involved in production.
Peru, like other developing countries, has
experienced strong urban growth during
the last few decades. Today, 72% of its 25
million inhabitants live in cities. Metropolitan Lima alone makes up 30% of
Peru's total population (INEI, 2000). As
migration from mountain regions to coastal
urban centers continues, urban consumer
preferences increasingly determine the
production priorities of the domestic
agricultura! sector, e.g., where production
and processing takes place, what farm
strategies are implemented, and what
technologies are applied. The geographic
proximity between producers and consumers is particularly important for fresh milk
and its derivatives, because of their high
perishability and bulkiness. This paper
1
2
Methods
Secondary literature, farm surveys (Julca,
2000; Senz, 2000) and census data (INEI,
1995), and milk collection statistics were
used to characterize Lima's dairy sector.
Based on that, three typical dairy farms
were defined (in terms of herd size): small
(1 O cows), medium (35 cows), and large
(220 cows). A farm-household optimization
model (Bernet, 2000) was used to assess
current profitability levels and the impact
on profits and production driven by
potential production context changes (e.g.,
feed prices and qual ity, herd management,
milk prices, and production levels). The
445
0.40
0.35
Vi
(/)
2.
Sf
(j
c.
CD
0.30
0.25
()
0.20
Results
0.15
Table 1. Characterization of small, medium, and large farms in coastal Peru near Lima.
Farm type
Small
Medium
Size of herd
< 20
20 to 100
Milking technology
by hand
by hand/machine
Artificial insemination
no/(yes)
yes/(no)
Labor force
family
contracted
On-farm specialists
no
no
High-quality feed mix
no
no/{yes)
Own silage production
no
no
Milk production (l/lactation)
3650
6800
Body weight of cow (kg)
450
650
First insemination date (mo.)
20
17
Calving interval (mo.)
> 15
14
Mortality rates
high
low
Sources: Avni, 1996; Julca, 2000; Senz, 2000.
446
Large
> 100
machine
yes
contracted
yes
yes
yes/(no)
7800
650
17
14
low
447
us - $
....----------.
2CXX)
Meat
O Milk
1500
lnfrastructure
Livestock
fZ.I sanitation
500
lnc~
(JIB Capital
l
1CXXJ
Labor
ITT Fodder (corn
t:;J with cobs)
EEI Feed
~ concentrate
Costs
Small
Medium Large
(10 cows) (35 cows) (220 cows)
448
Conclusions
Promising farm strategies
Milk prices in Lima will tend to fall when
the Lima market reaches a certain level of
saturation for bulky and perishable dairy
products. Therefore, the most important
strategy for farmers is to increase herd size
to benefit from better economies of size.
Small farm owners can simulate increased
herd size by forming local producer
associations linked to a milk collection
center to strengthen their negotiation
position vis-a-vis milk buyers and feed
providers. Such associations would provide
better access to production-relevant
information and technologies and could
faci litate other relevant activities for its
members (e.g., collective purchase and
References
Avni, S. 1996. La problemtica lechera en
el Peru. Estado de Israel - Fongal Lima
- Agencia Norteamericana para el
Desarrollo Internacional, Lima, Peru.
Bernet, T. 2000. The Peruvian dairy sector:
Farmer perspectives, development
strategies and policy options. PhD
Thesis. ETH No 13830, Zrich,
Switzerland.
INEI (Instituto Nacional de Estadstica e
Informtica). 1995. Censo agropecuario
1993. INEI, Lima, Peru.
INEI. 2000. Demographic indicators of
Peru in 1998 and 1999. Instituto
Nacional de Estadstica e Informtica,
Internet information (http://
www.inei.gob.pe/).
Julca, J. 2000. Caracterizacin productiva
de pequeos ganaderos lecheros del
Valle de Lurn. Thesis, Facultad de
Zootecnia, Universidad Agraria de La
Molina (UNALM), Lima, Peru.
McBride, E. 1997. Poltica alimentaria e
importacin de alimentos de origen
animal en el Per. Thesis, Facultad de
Zootecnia, Universidad Agraria de La
Molina (UNALM), Lima, Peru.
Senz, J. 2000. Uso de un modelo de
simulacin para la optimizacin de la
ganadera lechera en la zona de Lima.
Thesis, Facultad de Zootecnia,
Universidad Agraria de La Molina
(UNALM), Lima, Peru:
USDA. 1999. Peru: Dairy situation 1999.
USDA, Foreign Agricultura! Service,
Gain Report #PE9017, U.S. Embassy,
Lima, Peru.
449
CIP-Hanoi, Vietnam.
lnstitute of Ecology and Biological Resources (IEBR), Hanoi,
Vietnam.
451
452
30000
DProcessor
Non-processor
20000
15000
10000
Crops
Starch
Pig
101
74
70
35
280
25000
Total
Other
Total
453
Table 3. Perceptions of processors and non-processors toward the wastewater and salid waste generated
by starch processing (% of households responded "yes").
Processing villages
Non-processing
Questions
Processor Non-processor
villa ge
(N=142)
(N=103)
(N=35)
22
7
Ponds and lakes can still be used as clean water far
washing and cleaning.
77
78
87
Ponds and lakes are dirtier in processing season.
01
Ponds and lakes in processing villages are dirtier than non100
99
processing village.
51
48
o
Noticed plants nearby waste water ditches and canals have
died.
21
28
Saw children unintentionally swimming in wastewater.
58
23
28
20
Saw children get diseases or illness from falling into
wastewater.
o
Salid waste looks bad and dirty.
89
100
o
100
93
Noticed bad smell from salid waste.
84
98
63
Decomposed salid waste has negative effects on human's
health.
o
Noticed processors dumping salid waste in front of other
92
98
peoples' houses.
o
85
Noticed such dumping behavior cause conflict among
53
peo ple.
1 Oin this column indicates question was not relevant to non-processing villagers.
Table 5. Characteristics of starch-processing waste water collected at the point of processing, general
industrial waste water, and critica! values far regulatory compliance (mg/I).
Parameters
Parameter concentrations from various waste waters
Critica! value 1
Waste water at
Waste water in
General
Allowed Allowed
processing (mg/I)
canals (mg/I)
industrial
Value 2 Value 3
Cassava
Canna
Processing Non-pro. waste water
villages villages
(mg/I)
pH
4.6
5.8
6.9
7.55
6.85
5-9
5.5 - 9
BOD5
551.5
486.8
710
25
710
<50
<100
COD
6214.8
7378.8
1162
45.7
1162
<400
<100
SS
1466
3012.6
501
35
501
<100
<200
os
5735.7
7385.8
1.34
0.51
1340
Total P
34.82
72.37
24.21
25
24.21
<6
<8
Total N
168.1
199.3
102.5
12.87
102.5
<60
<60
Am-N
47.52
35.19
98.84
<1
<10
Mn
0.44
2.21
1.36
0.5<
1.36
<1
<5
so 4216
69.25
30
CN0.035
0.001
0.01
<0.1
<0.2
Note: 8005: biological oxygen demand; COD: chemical oxygen demand; SS: suspended solids; OS: dissolved solids;
Am-N: ammonium nitrogen; Mn: manganese; sol-: suttate ion; CN-: cyanide ion.
1 Based on Vietnam Wastewater Discharge Standards (MOSTE, 1995), Vietnam's standard No 5945 - 1995.
2 Allowed to be discharged only into water bodies used for navigation, irrigation, or aquatic breeding and cultivation.
(TCVN 5945-1995).
3 Allowed to be discharged only into the specific water bodies permitted by authoritative agencies.
455
Canna roots, with a sinuous and nonuniform shape, create much solid waste
during processing-comprising root skin
and fibrous residue. Cleaning canna roots
creates flaky solids and root skin accounting for about 5% of root weight. Solid
waste created during the first starch
separation stage consists of fiber, starch,
nutrients, and about 80-90% water.
Because canna starch is difficult to
extract, this residue still has a high starch
content, so it undergoes another manual
separation. After the second separation,
another household may take the solid
waste and put it through yet one more
cycle of separation to extract the very
small amount of starch remaining. Altogether about 5-7 kg of starch can be
extracted from one t of waste. The processors are willing to take the pain to extract
canna starch three times because canna
starch and noodles command much higher
prices and provide higher profits than does
cassava starch.
Laboratory analysis showed that cassava
solid waste contains 87.5% moisture and
12.5% dry matter. The dry matter is 69%
starch, 0.4% NPK, and 30% fiber, and
hence makes good pig and fish feed. Of
the 35,000 t of cassava solid waste
generated in 1999-2000, processors fed
11 % to their own pigs, and sold 71 % as
raw material for feed; only 18% was
wasted. Can na sol id waste, on the other
hand, contains less moisture (about 80%)
and more dry matter (20%), but the dry
matter contains only 4% starch and 0.6%
NPK, the rest being fiber that pigs cannot
easily digest. Canna solid waste is therefore hardly ever used as pig feed, and 90%
of it is abandoned to decompose or
dumped into lakes and ponds. In 19992000 about 15,000 t of sol id canna waste
was disposed of in this way. However, in
2001 a new model canna-starch separator
was introduced into the processing villages. lt has greater extracting efficiency
and creates extremely fine residues that
simply float away with the waste water, so
the amount of canna waste was greatly
reduced and little was abandoned in the
456
Conclusions
In the past, villagers caught fish and
mussels in the waterways near their
vil lages, but toda y there are no fish left.
As a result of processing activities, the
major canal that flows through the three
processing villages and past sorne nonprocessing ones has been filled with
mud and waste for several years. Sorne
sections of the canal are now only 0.5 m
deep. During the heavy rains between July
and August, roads in the processing
villages are now frequently flooded up to
0.2-0.3 m deep.
The decomposing or decomposed abandoned solid wastes, combined with
intensive pig-raising as an associated
enterprise, create foul odors that pollute
the processing and non-processing villages
alike. Sorne villages have set aside areas
for solid waste dumping, but the quantities
of wastes generated by processing have so
far overwhelmed this limited infrastructure.
Now that the extent, characteristics, and
impacts of the waste water and solid waste
References
Demo-os, R.A., T.S.J. Valdez, and M.C.
Mapili, Jr. 2000. Feeding value of
protein-enriched sweet patato pulp for
broilers. Philippines Journal of Veteran
Animal Science 26:41-50.
MOSTE (Ministry of Science, Technology,
and Envioronment). 1995. Vietnamese
government standard of environment.
Hanoi Publishing House. Hanoi,
Vietnam.
Pham, C.B., M.R.L.Y. Lat, T.J. Ramirez,
M.J. Quinlat, and L.J. Pham. 1992.
Enriching cassava protein using solid
state fermentation. BIOTECH,
University of the Philippines at Los
Banas, Philippines.
UNEP (United Nations Environmental
Program). 1994. Overview of
environmental indicators: State of the
art and perspectives. IVM, Netherlands.
457
d
DAS
DAT
DADO
DAN IDA
DEM
DFID
DM
DNA
Bacillus thuringiensis
blight unit
bacteria! wilt
cellu lose acetate electrophoresis
cleaved amplified polymorphic sequence
Cation exchange capacity
Civil Engineers Network Africa, South Africa
Centro lnternacinal de Agricultura Tropical (Colombia)
Centro lnternacinal de Investigaciones para el Desarrollo (IRDC)
lnternational Maize and Wheat lmprovement Center (Mexico)
lnternational Potato Centro (Centro Internacional de la Papa)
centimeter
cassava mosaic virus disease
cubic feet per minute
Consortium for the Sustainable Development of the Andean
Ecoregion
Central Potato Research lnstitute (India)
Centre for Plant Breeding and Reproduction Research (Netherlands)
Central Tuber Crops Research lnstitute, lndia/lndian Council of
Agricultura! Research
day
days after sowi ng
days after transplanting
District Agriculture Development Office (Nepal)
Royal Danish Ministry of Foreign Affairs
digital elevation model
Department for lnternational Development
dry matter
deoxyribonucleic acid
459
EARO
EARRNET
EC
ECABREN
ELISA
EPA
ESARC
FAO
FAPESP
FFS
FMV
FORTIPAPA
FPR
FSP
FU
GCA
GILB
GIS
GOES
Gpi
GPS
GTZ
GUS
GxE
h2
HDP
HPAEC
HYV
IAF
IAl-ISP
IARC
ICM
ICRAF
ICRISAT
IDM
IDRC
IFAD
IFAS
IFDC
IFPRI
llRDEP
llTA
ILRI
INERA
460
INFOANDINA
INIA
INIAP
INRA
INTA
IOFM
IPGRI
IPM
IRR
IRRI
ISO
J
K
KARI
kg
1
LB
LEACHM
LMF
m
mm
masl
MCL
mo
MRR
n
N/m2
NAFTA
NAR
NARO
NARS
NASH
NCM
NDVI
NGO
NPV
NUE
OFE
OM
OPEC
p
p
PAGE
PapMV-U
PARC
ppb
PBS
PCA
PCR
pep
461
PGI
pH
PI
PIC
PLRV
PNS-PRODISE
PO
PoGV
PRAPACE
PRECODEPA
PRGA
PROINPA
PRONAMACHCS
PSTVd
PTL
PTM
PVT
PVX
PVY
QTL
RAPO
rAUDPC
RCBD
rDNA
RFLP
RGR
RH
RKN
RRA
RRD
RS
SAAS
SAPPRAD
SARDl-UMCOR
SARRNET
SCA
SCRI
SDC
SDRF
sos
SEINPA
SE NASA
SG
462
SIUPA
SLA
SM-CRSP
SPCFV
SPCSV
SPFMV
SPLV
SPVD
SQR
SSR
t
TAC
TCRC
TPS
UAP
UMMV
UMV
UNEP
UNHEVAL
UNICEF
UPGMA
UPWARD
USAID
uv
uve
w
WEPP
WUE
463
Author lndex
Alarcn, Lidia. Crop lmprovement and
Genetic Resources, lnternational Potato
Center, A.P. 1558, Lima 12, Peru. p. 95.
Aley, Pedro. Crop Protection, lnternational
Potato Center, A.P. 1558, Lima 12, Peru.
p. 105.
Amaro, J.R. Universidad Nacional del
Centro del Peru, Huancayo, Peru. p. 69.
Ames, Teresa. Crop Protection,
lnternational Potato Center, A.P. 1558,
Lima 12, Peru. p. 397.
Amoros, Walter. Crop lmprovement and
Genetic Resources, lnternational Potato
Center, A.P. 1558, Lima 12, Peru.
p. 185.
Asmunati, Rini. Mitra Tani, Yogyakarta,
Indonesia. p. 331.
Baigorria, Guillermo. Production Systems
and Natural Resource Management,
lnternational Potato Center, A.P. 1558,
Lima 12, Peru. p. 361, 371.
Barrea, O. Fundacin para la Promocin e
Investigacin de Productos Andinos
(PROINPA), 191 Calle Honduras, Barrio
Petrolero, Sucre, Bolivia. p. 143.
Barreda, Carolina. Production Systems and
Natural Resource Management, A.P.
1558, lnternational Potato Center, Lima
12, Peru. p. 361.
Bejarano, Carlos. PROINPA Foundation,
Cochabamba, Bolivia. p. 251.
Bello, Vernica. Crop lmprovement and
Genetic Resources, lnternational Patato
Center, A.P. 1558, Lima 12, Peru.
p. 267.
Beltrn, Gregario. Crop lmprovement and
Genetic Resources. lnternational Potato
Center, A.P. 1558, Lima 12, Peru. p. 63.
Bernet, Thomas. Social Science,
lnternational Potato Center, A.P. 1558,
Lima 12, Peru. p. 445.
Bi, Y.P. Shandong Academy for
Agricultura! Sciences, Jinan, Shandong,
China. p. 219.
Boncodin, Raul. CIP-UPWARD, PCARRD
Complex, Los Banos, Laguna,
Philippines. p. 433.
465
466
Author lndex
p. 211.
Garrett, Karen A. Kansas State University,
Manhattan, KS, USA. p. 225.
Garry, Guillemette. CIP/Present address:
c/o 7 rue de Concise, 53940
St. Berthevin, France. p. 39.
Gastelo, Manuel. Crop lmprovement and
Genetic Resources, lnternational Patato
Center, A.P. 1558, Lima 12, Peru. p. 63.
Gaur, P.C. Central Patato Research
lnstitute (CPRl/ICAR), Shimla, H.P.,
India. p. 219.
Ghislain, Marc. Crop lmprovement and
Genetic Resources, lnternational Potato
Center, A.P. 1558, Lima 12, Peru. p. 27,
167, 175, 267, 295.
Golmirzaie, Ali M. Department of
Horticulture, University of Arkansas,
Fayetteville, AR 72701, USA. p. 175.
Gomez, Carlos. Universidad Nacional
Agraria, La Molina, Lima 12, Peru.
p. 445.
Gomez, Rene. Crop lmprovement and
Genetic Resources, lnternational Potato
Center, A.P. 1558, Lima 12, Peru.
p. 175.
Gonzles, Eliana. Crop Protection,
lnternational Patato Center, A.P. 1558,
Lima 12, Peru. p. 39.
Gonzales, Silvia. PROINPA Foundation,
Cochabamba, Bolivia. p. 251.
Grande, Enrique. Crop lmprovement and
Genetic Resources, lnternational Patato
Center, A.P. 1558, Lima 12, Peru.
p. 219.
467
468
Author lndex
p. 105.
Salazar, luis F. Crop Protection,
lnternational Potato Center, A.P. 1558,
lima 12, Peru. p. 267.
Sanchez, Pablo A. Asociacin para el
Desarrollo Rural de Cajamarca
(ASPADERUC), Cajamarca, Peru.
p. 413.
Santa Cruz, Magnolia. Crop lmprovement,
lnternational Potato Center, A.P. 1558,
Lima 12, Peru. p. 39, 381.
Sattelmacher, Burkhard. lnstitute of Plant
Nutrition and Soil Science, Kiel
University, Germany. p. 273.
Simon, Reinhart. Crop lmprovement and
Genetic Resources, lnternational Potato
Center, A.P. 1558, lima 12, Peru. p. 27.
469
470
Author lndex
471
472
p. 55-72.
Fuglie, K.O., C. Narrod, and
C. Neumeyer. 2000. Public and private
investments in animal research. In:
Fuglie K.O. and O.E. Schimmelpfennig
(eds.). Public-private collaboration in
agricultura! research. lowa State
University Press, Ames, IA, USA.
p. 117-151.
Fuglie, K.O. 1999. Conservation tillage
and pesticide use in the Cornbelt.
Journal of Agricultura! and Applied
Economics 31:133-147.
Fuglie, K.O. 2000. Trends in agricultura!
research expenditures in the United
States. In: Fuglie, K.O. and O.E.
Schimmelpfennig (eds.). Public-private
collaboration in agricultura! research.
lowa State University Press, Ames, IA,
USA. p. 9-23.
39:131-148.
Fuglie, K. 1999. lnvesting in agricultura!
productivity in Indonesia. Forum
Penelitian Agro Ekonomi 1 7: 1-16.
46:547-555.
Ghislain, M., M. Bonierbale, and
R. Nelson. 1999. Gene technology for
potato in developing countries. In:
Hohn, T. and K.M. Leisinger (eds.).
24 p.
He, W., Z.M. Zhang, and Y. Wang. 1999.
GILB meeting in Ecuador and late blight
research progress. Chinese Potato
Journal 13:182-183.
473
georeferenciada de la distribucin
global de la papa. Sistmica 1 :19-24.
474
p. 1-13.
Prain, G. and
Prain, G.,
475
476
477
Networks
CONDESAN (Consortium for the Sustainable Development of the Andean
Ecoregion)
(same address, telephone, and fax as CIP
headquarters)
E-mail: condesan@cgiar.org
Website: www.condesan.org
Contact: Elias Mujica, Acting Coordinator
GILB (Global lnitiative on Late Blight)
(same address, telephone, and fax as CIP
headquarters)
E-mail: gilb@cgiar.org
Website: www.cipotato.org/gilb
Contact: Wanda Collins, GILB Coordinator
SUB-SAHARANAFRICA(ssA)
478
Networks
PRAPACE (Regional Potato and
Sweetpotato lmprovement Program for East
and Central Africa)
479
Sol 4 825
Nam Thanh Cong
Lang Ha, Dong Da
Hanoi, Vietnam
Tel and fax: +84 4 835 5494
E-mail: cip-hanoi@fpt.vn or
d.peters@cgiar.org or dpeters@fpt.vn
Contact: Dai Peters, Postharvest Specialist
Networks
ANSWER (Asian Network for Sweetpotato
Genetic Resources)
c/o CIP-ESEAP Regional Office
JI. Raya Ciapus
Bogor, Indonesia
480
Telephone: 62-251-317951
Fax: 62-251-316264
E-mail: CIP-Bogor@CGIAR.Org or
S.Mahalaya@CGIAR.Org
Website: www.eseap.cipotato.org/answer/
Contacts: Algerico M. Mariscal, ANSWER
coordinator
* Central and Eastern Europe, Transcaucasia, and Central Asia (ECA): closed
in 2000