Animal Reproduction Science 6061 2000.

605614
www.elsevier.comrlocateranireprosci

Epidemiology of reproductive performance in dairy

cows
Y.T. Grohn
a,) , P.J. Rajala-Schultz a,b
a

Section of Epidemiology, Department of Population Medicine and Diagnostic Sciences, College of Veterinary
Medicine, Cornell Uniersity, Ithaca, NY 14853 ,USA
b
Department of Veterinary Preentie Medicine, College of Veterinary Medicine, The Ohio State Uniersity,
Columbus, OH 43210, USA

Abstract
The objectives of this presentation are to review results of our previous and on-going research
with respect to the risk factors and consequences of poor reproductive performance in dairy cows,
and to develop an economic framework to optimize decisions related to dairy cow reproductive
performance. To make profitable breeding and replacement decisions, the farmer must account for
factors including age, production level, lactation stage, pregnancy status, and disease history of the
cows in the herd. Establishing the interrelationships among disease, milk yield, reproduction, and
herd management is necessary for developing a decision model for disease treatment, insemination, and replacement.
The data for the studies reviewed in this presentation incorporate health, production, and
management components from Holsteins in the Northeast USA and Ayrshires from Finland. Data
were analyzed using the Cornell Theory Center Supercomputer. The effect of risk factors on
reproductive disorders was modeled with logistic regression, and on conception, insemination, and
culling with survival analysis. The effect of reproductive disorders on milk yield was analyzed
with mixed models. Economic optimization of reproductive performance was done with dynamic
programming DP..
High milk yield, high parity, and calving in winter were risk factors for several reproductive
disorders. These disorders, in turn, delayed insemination and conception in dairy cows, and some
of them increased the risk of culling. Dystocia, retained placenta, and early metritis led to a
short-term drop in milk production. High milk yield was not a major factor in delaying
conception, except in first parity cows. However, higher yielders were more likely to be
inseminated, and less likely to be culled. Non-pregnant cows had a higher risk of being culled.

Corresponding author. Tel.: q1-607-253-3571; fax: q1-607-253-3083.

E-mail address: ytg1@cornell.edu Y.T. Grohn
..

0378-4320r00r$ - see front matter q 2000 Elsevier Science B.V. All rights reserved.
PII: S 0 3 7 8 - 4 3 2 0 0 0 . 0 0 0 8 5 - 3

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Y.T. Grohn,
P.J. Rajala-Schultzr Animal Reproduction Science 6061 (2000) 605614

Reproductive performance of dairy cows influenced a herds profitability, and good heat
detection and conception rates provided opportunities for management control. It was not always
economically advantageous to get cows pregnant as soon as possible, and there was no one
optimal value for the calving interval length for all cows in a herd. q 2000 Elsevier Science B.V.
All rights reserved.
Keywords: Dairy cows; Reproduction; Epidemiology

1. Introduction
Profitable milk production relies upon a careful, efficient and cost-effective management of dairy herds. Poor reproductive performance is an important production-limiting
factor. Management decisions on dairy farms should be economically justified. Our
ultimate goal is to help farmers make the best decisions regarding management of their
dairy herds. To do this, we must understand the biological parameters involved, namely
milk yield, disease, and reproductive performance.
Although many studies on reproductive performance in dairy cows have been
conducted, few have integrated together epidemiology and economics. Many of them
have been constrained by the lack of three factors: a large health database, computing
power, and sophisticated statistical tools. Fortunately, we have had access to high quality
data on Finnish Ayrshires beginning with Grohn
et al. 1986., and continuing until the
.
present ; such data have not been previously available in the United States, where
Holsteins predominate. Therefore, we established a health and production database in
the Northeast USA beginning with Grohn
et al. 1995., and continuing until the
.
present to study the epidemiology and economics of production diseases under Northeastern management.
In this presentation, our focus is reproductive performance in dairy cows. We will
concentrate on three aspects of it: 1. risk factors for poor reproductive performance; 2.
consequences of poor reproductive performance; and 3. economic optimization of
reproductive performance. The material for this review was primarily taken from our
on-going epidemiological research on New York Holsteins and Finnish Ayrshires.

2. Risk factors for poor reproductive performance

2.1. Increased risk of reproductie disorders
The dairy industrys major goal is to provide milk for the consumer market. Over the
past several decades, milk yield of cows has increased markedly. However, one negative
impact of this improvement is reduced fertility e.g., cows open for longer, more
services per conception. and higher incidence of reproductive problems, which, in turn,
contribute to reduced fertility e.g., Dhaliwal et al. 1996... Some have argued that
higher disease incidence and reduced fertility in high-producing herds result more from
poor management than from biology. High milk yield may contribute to a negative

Y.T. Grohn,
P.J. Rajala-Schultzr Animal Reproduction Science 6061 (2000) 605614

607

energy balance in some cows, especially those still growing, and disease may occur.
Selective culling muddies the issue further. High yielders are more likely to remain in
the herd, and receive more veterinary treatment, even when they become ill, than low
yielders.
To address this question, we conducted several studies in both Finnish Ayrshires and
New York Holsteins e.g., Grohn
et al., 1990, 1994, 1995; Rajala and Grohn,
1998a..
Lactational incidence risks for reproductive disorders in our latest Finnish study Rajala
and Grohn,
1998a. were: dystocia, 2.1%; retained placenta, 3.1%; metritis, 3.2%; silent
heat, 8.1%; ovarian cysts, 7.3%; and other infertility, 1.9%. In our New York study
Grohn
et al., 1995., incidences were higher: retained placenta, 7.4%; metritis, 7.6%;
and ovarian cysts, 9.1%.
Using logistic regression, we analyzed the effect of several risk factors on the
occurrence of reproductive disorders. In one study Grohn
et al., 1990., first parity cows
were most likely to have dystocia Table 1.. Older cows were more likely to have
retained placenta and ovarian cysts. Winter calvers had higher risks of early metritis,
silent heat, ovarian cysts, and other infertility than summer calvers did. Most reproductive disorders were interrelated. For example, dystocia was a risk factor for retained

Table 1
Risk factors parity, calving season, diseases. for seven diseases, in 61,124 Finnish Ayrshires. All seven
models included community as a proxy of management a
Risk factors

Diseases
DystociaRetained placentaOvarian CystsEarly metritisLate metritisSilent heatOther infertility

Parity
1
2
3q

2.1
1.0
U
1.3

1.0
1.5U

1.0
U
1.2
U
1.4

Caling season
JanApr
MayAug
SeptDec

1.0
1.0
U
1.6

1.6
1.0
U
1.7

4.1U
U
19.8

U
1.5
U
2.9
U
1.5
U
3.3

U
2.8

3.2
U
3.7
U
4.4

U
5.4

Diseases
Dystocia

Abortion

Retained placenta
Early metritis

Late metritis

Silent heat

Prolapsed vagina
Ovarian cyst

Other infertility
a

U
1.6
1.1

1.0
U
1.6
U
1.9

1.9
U
1.9
1.0

1.3
1.3
1.0

U
3.5
U
2.5
U
1.6

U
2.2
U
3.6

U
2.1
U
1.3
U
1.5
U
3.0

U
1.6

U
1.7
U
2.4

U
2.6

U
1.7
1.1

Values are odds ratios, which measure how much more or less likely the outcome is among observations
with a given risk factor or level of risk factor if ) 2 categories., compared with those without it or reference
category, which has an odds ratio of 1.0.. Milk yield was also included in these models.
U
p- 0.05.

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Y.T. Grohn,
P.J. Rajala-Schultzr Animal Reproduction Science 6061 (2000) 605614

placenta and metritis. Results were similar in our study on New York Holsteins, for
parity and calving season, but there were fewer disease associations Grohn
et al., 1995..
Cows with higher milk yield in the previous lactation were more likely to have
retained placenta, early metritis, silent heat, ovarian cysts, and other infertility than were
cows with lower milk yield. Cows with higher milk yield in the current lactation were
more likely to have ovarian cysts than were cows with lower milk yield. In a similar
study on New York Holsteins Grohn
et al., 1995., higher milk yield was associated
only with ovarian cysts.
In addition to breed difference, other factors may account for the greater number of
disorders associated with milk yield in Ayrshires than Holsteins. Finnish farms are much
smaller than New York farms so management differs. The Finnish studies contained
over five times as many cows as our New York study, so smaller differences become
significant. Also, in New York, a farmer may treat animals for disease, but in Finland
only veterinarians treat animals.
2.2. Risk factors for delayed conception
Many factors influence reproductive performance in dairy cows, only some of which
are under the farmers control. Milk yield may play an important role in reproduction.
However, the relationship between milk yield and conception is difficult to study
because of the confounding effect of culling. Conception does not necessarily occur at
the first breeding; cows may have to be inseminated several times. A cow that remains
open i.e., does not conceive. for a long time or has low milk yield is more likely to be
culled. Some researchers have argued that high yielders are less fertile than low yielders
are; this association may, however, be complicated by selective culling.
The objective of our New York study was to measure the effect of 60-day cumulative
milk yield on rates of conception and first breeding in lactation Eicker et al., 1996..
Data were from 15,320 Holsteins calving between June 1990 and November 1993, in 26
herds. We used survival analysis, which allows inclusion of all cows, whether or not
they had conceived or been bred by the end of the study. Thus, the loss of information
was minimized. Parity, calving season, and herd as a proxy of management were
included as confounders.
Retained placenta, metritis, and ovarian cysts were risk factors for conception Table
2.. Cows with retained placenta had a 14%, those with metritis a 15%, and those with
ovarian cysts a 21% lower conception rate than cows free of these disorders. Current
cumulative 60-day milk yield had no effect on conception rates, except among the
highest yielders. They had a slightly lower but not significant. conception rate than did
the lowest yielders.
Because these results may be a consequence of farmers intentionally delaying
insemination of high yielding cows, we also studied the association between milk yield
and time to first insemination. As 60-day milk yield increased, so did insemination rates.
The highest yielders were nearly 30% more likely to be inseminated than were the
lowest yielders.
Our results on New York Holsteins indicate that milk yield in the first 60 days has
only a minimal effect on conception. Only the highest yielders had a slightly lower

Y.T. Grohn,
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609

Table 2
Effect of milk yield, parity, calving season, and disease on conception in 13,307 New York Holsteins
Risk factor

Hazard ratio a

First 60-day cumulatie milk yield (kg)

F1582
15831891
18922195
21962541
) 2541

1.0
0.99
1.01
1.01
0.92

Parity
1
2
G3

1.0
0.98
UU
0.92

Caling season
DecFeb
MarMay
JunAug
SepNov

1.0
U
0.93
1.06
1.01

Disease
Retained Placenta
Metritis
Ovarian Cysts

0.86
UU
0.85
UU
0.79

UU

a
Hazard ratios for factors in proportional hazards model. The hazard ratio is the ratio between two relative
risks of an event e.g., conception.. For example, if a cow has a hazard ratio of 0.92 parity G 3., then she has
an 8% reduced likelihood of conceiving than a first parity cow.
U
p- 0.05.
UU
p- 0.01.

conception rate than their herdmates. Older cows and sick cows were less likely to
conceive. In contrast, the rate of being bred increased with 60-day milk. These findings
demonstrate that farmers are making rational decisions by breeding young, healthy, high
yielding cows.
Among 30,036 multiparous Finnish Ayrshires, the lowest yielders were less likely to
conceive than were average yielders Harman et al.,1996a.. Among 11,761 heifers, the
highest yielders were less likely to conceive than were average yielders. A number of
disorders decreased conception probability in both multiparous and primiparous cows:
anestrus, ovulatory dysfunction, other infertility, late metritis, and clinical ketosis
Harman et al., 1996b..

3. Consequences of poor reproductive performance

3.1. Lowered milk yield due to reproductie disorders
It seems obvious that disease causes milk loss, but surprisingly, some studies have
found that increased milk yield is associated with disease. How milk yield is expressed

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Y.T. Grohn,
P.J. Rajala-Schultzr Animal Reproduction Science 6061 (2000) 605614

in the analysis is important; a single summary measure of milk yield e.g., 305-day
yield. may give a completely different answer than monthly milk yields Detilleux et al.,
1994..
In one of our studies Rajala and Grohn,
1998b., we estimated the effects of dystocia,
retained placenta, and metritis on milk yield using repeated, monthly test day milk yields
taken on 37,776 Finnish Ayrshire cows. All three disorders had a significant effect on
milk yield. Their impact varied across parities. Milk losses were greatest immediately
following diagnosis. For instance, in parity 2, daily losses from dystocia, retained
placenta, and early metritis during the 2 weeks following diagnosis were 2.2, 1.4, and
1.3 kgrday, respectively.
There will probably always be some loss of milk yield after a disease. It is important
to use proper methodology when quantifying the loss, i.e., it is more accurate to use
monthly measurements of milk yield rather than a single, summary lactational measure.
Only in this way can true losses be observed. The measurement of loss of milk yield
following disease is important. Knowing whether the milk loss due to disease is small
and temporary or larger and more sustained helps a farmer decide whether it is worth
keeping the cow in the milking herd.
3.2. Shortened productie life
Culling is a complex issue: many factors are involved. Dairy cows are culled for
involuntary i.e., death, acute disease and infertility. and voluntary i.e., low yield.
reasons. Both biology and management affect the culling decision. Reproductive failure
is a major reason for involuntary culling Fetrow, 1988.. However, when making a
decision, the dairy farmer considers at least five major reasons: parity, milk yield, stage
of lactation, diseases, and conception status.
Using survival analysis with time-dependent covariates, we studied the effect of
seven diseases on culling in New York Holsteins. The effects of milk yield and
conception status on culling were also studied, and the interactions between these
covariates and stage of lactation were accounted for Grohn
et al., 1998.. Cows that had
not yet conceived, as well as older cows and cows with certain diseases, had a higher
risk of culling. For instance, open cows i.e., before conception. were seven-and-a-half
times more likely to be culled than pregnant cows.
We have also conducted a series of studies on Finnish Ayrshires, of increasing
complexity, starting with only disease effects on culling in the model, then adding
pregnancy status effects, and finally adding milk yield effects Rajala-Schultz and
Grohn,
1999a,b,c.. All models also included parity, calving season, and herd as

covariates. Pregnancy status was the most influential factor affecting culling decisions,
followed by milk yield. Several diseases also significantly affected culling. The effects
of all these factors varied with the stage of lactation. Dystocia and metritis increased the
risk of culling at the time of their occurrence, but also at the end of lactation. Ovarian
cysts and anestrus also affected culling decisions. For instance, if a cow had been
diagnosed with ovarian cysts or anestrus within 150 days after calving, she was less
likely to be culled than a cow without these disorders. Here, it is important to remember
that in Finland diagnoses are made by veterinarians and by calling a veterinarian to a
farm, the farmer has already made a preliminary decision to treat and keep the cow. Of

Y.T. Grohn,
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611

Table 3
Effects of pregnancy status and number of inseminations on culling in 39,727 Finnish Ayrshire cows calving
in 1993. The model also included parity, season, and herd
Pregnancy status
Lactational stage of culling day.

Lactational stage of conception day. a

Risk ratio

0150

0150
0
0150
151240
0
0150
151240
241305
0
0150
151240
241305
) 305
0

1.0
UUU
25.8
1.0
1.3UUU
1.6UUU
1.0
2.1UUU
2.6UUU
4.9UUU
1.0
1.7UUU
2.9UUU
3.8UUU
UUU
23.9

151240

241305

) 305

Number of inseminations
0
1
2
3
)3

UUU

10.0
1.0
0.9UUU
0.9U
0.9UU

a
Stage of lactation when the farmer was assumed to know when a cow was pregnant; 0 refers to open cows
i.e., cows that did not conceive at all..
U
p- 0.05.
UU
p- 0.01.
UUU
p- 0.001.

the 15 diseases studied, retained placenta was the only disorder without an effect on
culling.
Effects of pregnancy status and number of inseminations on culling of Finnish
Ayrshires are in Table 3. The data are expressed as risk ratios, which measure the risk of
a cow being culled depending on the time when she was known to be pregnant
compared with a reference level i.e., pregnant by day 150 after calving, RR s 1.0.. The
knowledge about a cows pregnancy status had a different effect on culling at different
stages of lactation. The later the cow conceived, the higher was the risk of her being
culled. Cows not inseminated at all were at 10 times higher risk of being culled than
cows inseminated only once. Cows inseminated more than once were slightly less likely
to be culled than cows inseminated only once.
4. Economic optimization of reproductive performance
Reproductive performance of a dairy herd affects profitability of the farm. Reproductive inefficiency reduces milk yield and number of calves born i.e., replacements

612

Y.T. Grohn,
P.J. Rajala-Schultzr Animal Reproduction Science 6061 (2000) 605614

produced., and may increase the cost of veterinary services and, as shown above, it also
affects the culling rate. Insemination, treatment, and culling decisions represent an area
of dairy herd management affecting profitability. To make rational decisions, the farmer
needs a valid estimate of the future profitability of each cow, accounting for factors
including age, production level, lactation stage, pregnancy status, and disease history.
Dynamic programming DP. can be used for optimizing a sequence of interrelated
decisions. The objective of DP models developed for optimizing breeding and replacement decisions in dairy herds is to maximize the expected net present values from
current and replacement cows over a given decision horizon see, e.g., DeLorenzo et al.
1992... Based on net present values, decisions are made monthly to either 1. keep and
inseminate, 2. keep and not inseminate, or 3. replace a cow. If the net present value of
a cow at a certain stage is lower than that of a replacement heifer, a decision to replace
her should be made. Similarly, if the net present value from a cow leaving her open is
lower than breeding her immediately, a decision should be made to breed her.
We used a DP model, described by McCullough 1992., to determine the structure of
an optimal herd under Finnish conditions Rajala-Schultz et al., 1999a.. The calving
interval resulting from the optimal replacement and breeding decisions over a 5-year
decision horizon was 363 days; however, there was considerable seasonal variation
within parities, depending on the month of calving. It was optimal to have most calvings
in the fall, and to keep the calving interval of these cows approximately 1 year. We also
estimated the effects of reproductive performance on optimal breeding decisions and
profitability of Finnish dairy herds Rajala-Schultz et al., 1999b.. It was not optimal to
start breeding cows calving in spring and early summer immediately after the voluntary
waiting period; instead, it was preferable to allow the calving interval to lengthen for
these cows so that their next calving was in the fall. However, cows calving in the fall
should be bred immediately after the voluntary waiting period. These findings were
mainly due to seasonal milk pricing and higher milk production among fall calvers in
Finland. Reproductive performance had a considerable impact on a herds profitability;
good heat detection and conception rates provided opportunities for management
control.

5. Conclusions
Reproductive performance affects dairy herd profitability by decreasing milk yield
and the number of calves produced, and by increasing the culling rate. Common sense
would dictate that to improve reproductive efficiency, the occurrence of reproductive
disorders must be minimized. To appreciate the complexity involved in measuring the
associations discussed in this presentation, one must realize the limits of epidemiological
research. The difficulty is that what is observed is the product of both biology and
environment. Selection bias also plays a role; cows are managed, bred, and culled
selectively. Nevertheless, with proper study design and control of confounding factors
like parity, calving season, and herd, valid and largely objective estimates can be
obtained.

Y.T. Grohn,
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613

Having quantified these biological parameters, they can be incorporated into an

economic model to see their impact on a farms profitability. The result of this DP
approach is the net present value of a cow; this model also provides an insemination
value for each cow in a herd, indicating how much could be spent in trying to get a cow
pregnant. This information helps farmers make rational decisions to maximize the
profitability of their dairy herds. Thus, seeing the farm as an enterprise where all
management decisions including breeding and treatment decisions. must be economically justified is a strategy for optimizing herd health and efficiency, and, ultimately, its
livelihood.

Acknowledgements
The initial work of this project was the AVMA Foundation Research Proposal The
effect of high milk production on disease and reproduction in dairy cows: logistic and
survival analysis, 19911992. Major funding for the project was provided by the
USDA National Research Initiative Competitive Grants Program award number 9203884., 19921995 and award number 94-02316., 19941997. The current research on
optimizing and delivering dairy herd health and management decisions is funded by the
USDA Rural Fund of America award number 97-05048., 19982000, and the USDA
National Research Initiative Competitive Grants Program award number 98-02405.,
19982001. This research was conducted using the resources of the Cornell Theory
Center Cornell University, Ithaca, NY.. The most recent research conducted on Finnish
Ayrshires was supported by the Finnish Academy.

References
DeLorenzo, M.A., Spreen, T.H., Bryan, G.R., Beede, D.K., van Arendonk, J.A.M., 1992. Optimization model:
insemination, replacement, seasonal production and cash flow. J. Dairy Sci. 75, 885896.
Detilleux, J.C., Grohn,
Y.T., Quaas, R.L., 1994. Effects of clinical ketosis on test day milk yields in Finnish
Ayrshire cattle. J. Dairy Sci. 77, 33163323.
Dhaliwal, G.S., Murray, R.D., Dobson, H., 1996. Effects of milk yield, and calving to first service interval, in
determining herd fertility in dairy cows. Anim. Reprod. Sci. 41 2., 109117.
Eicker, S.W., Grohn,
Y.T., Hertl, J.A., 1996. The association between cumulative milk yield and days open
and days to first breeding in New York State Holstein cows. J. Dairy Sci. 79, 235241.
Fetrow, J., 1988. Culling of dairy cows. In: 20th annual meeting of AABP, Phoenix, Arizona, AABP.
Grohn,
Y.T., Eicker, S.W., Ducrocq, V., Hertl, J.A., 1998. The effect of disease on culling Holstein dairy
cows in New York State. J. Dairy Sci. 81, 966978.
Grohn,
Y.T., Eicker, S.W., Hertl, J.A., 1995. The association between previous 305-day milk yield and disease
in New York State dairy cows. J. Dairy Sci. 78, 16931702.
Grohn,
Y.T., Erb, H.N., McCulloch, C.E., Saloniemi, H.S., 1990. Epidemiology of reproductive disorders in
dairy cattle: associations among host characteristics, disease and production. Prev. Vet. Med. 8, 2539.
Grohn,
Y.T., Hertl, J., Harman, J.L., 1994. The effect of early lactation milk yield on reproductive disorders in
dairy cows. Am. J. Vet. Res. 55, 15211528.
Grohn,
J., 1986. An epidemiological and genetic study on registered diseases
Y.T., Saloniemi, H., Syvajarvi,

in Finnish Ayrshire cattle: I. The data, disease occurrence and culling. Acta Vet. Scand. 27, 182195.

614

Y.T. Grohn,
P.J. Rajala-Schultzr Animal Reproduction Science 6061 (2000) 605614

Harman, J.L., Grohn,

Y.T., Erb, H.N., Casella, G., 1996a. Event-time analysis of the effect of 60-day milk
production on dairy cow interval-to-conception. Am. J. Vet. Res. 57, 634639.
Harman, J.L., Grohn,
Y.T., Erb, H.N., Casella, G., 1996b. Event-time analysis of the effect of calving season,
parity, and concurrent disease on interval-to-conception. Am. J. Vet. Res. 57, 640645.
McCullough, D.A., 1992. Effects of model specifications and exogenous variables on a stochastic dynamic
insemination and replacement model for dairy cattle. MS thesis, Univ. Florida, Dept. of Dairy Science,
Gainesville, pp. 168.
Rajala, P.J., Grohn,
Y.T., 1998a. Disease occurrence and risk factor analysis in Finnish Ayrshire cows. Acta
Vet. Scand. 39, 113.
Rajala, P.J., Grohn,
Y.T., 1998b. Effects of dystocia, retained placenta, and metritis on milk yield in Finnish
Ayrshire cows. J. Dairy Sci. 81, 31723181.
Rajala-Schultz, P.J., Grohn,
Y.T., 1999a. Culling of dairy cows: Part I. Effects of diseases on culling in

Finnish Ayrshire cows. Prev. Vet. Med. 41, 195208.

Rajala-Schultz, P.J., Grohn,
Y.T., 1999b. Culling of dairy cows: Part II. Effects of diseases and reproductive
performance on culling in Finnish Ayrshire cows. Prev. Vet. Med. 41, 279294.
Rajala-Schultz, P.J., Grohn,
Y.T., 1999c. Culling of dairy cows: Part III. Effects of disease, pregnancy, and
milk yield on culling in Finnish Ayrshire cows. Prev. Vet. Med. 41, 295309.
Rajala-Schultz, P.J., Grohn,
Y.T., Allore, H.G., 1999a. Optimizing replacement decisions on Finnish dairy
herds: price effects of calving pattern, herdlife and calving interval. Acta Vet. Scand., May 20. in press.
Rajala-Schultz, P.J., Grohn,
Y.T., Allore, H.G., 1999b. Optimizing breeding decisions for Finnish dairy herds:
effects of reproductive performance on profitability. Acta Vet. Scand., June 15. in press.