J 1601-5223 1966 tb02053 X

H E R E 55-20
MUTATIONS AND CROP IMPROVEMENT. VII. THE GENUS ORYZA L. (GRAMINEAE)

By A K E GUSTAFSSON and W A R GADD
INTERNATIONAL ATOMIC ENERGY AGENCY, VIENNA I, AND
THE ROYAL COLLEGE OF FORESTRY, STOCKHOLM
50,
SWEDEN
(Received April 9th, 1966)
N our articles on Mutations and Crop Improvement. I-VI. (Hereditas 1965) the authors have dealt with mutation problems in species within the five genera Lupinus, Ipomoea, Poa, Arachis and Avena. The species analysed reproduce by seed or by vegetative means, by sexual or apomictic processes, and are diploid or polyploid. In some further articles we shall take up for discussion five more species or species groups, uiz. Oryza spp., especially Oryza sativa (rice), Solanum tuberosum (potatoe), Triticum spp. (wheat), Vitis vinifera (vine) and Phaseolus spp. (beans). As mentioned in our introductory paper of the series (I) we are aware that mistakes will appear in our analyses. We ask scientists concerned, or other experts, to inform us not only of mistakes made but also of literature references pertinent to the subject but not considered in our reviews.
I. ORIGIN AND GENETICS OF RICE

1. General references
ALIM et al., 1962 (rice research in East Pakistan); BIANCHI, 1965 (genetics and breeding) ; CHANDRARATNA, 1964 (monograph on rice genetics and breeding) ; T. T. CHANG,1964 (genetics and cytogenetics) ; CHANGand BARDENAS, 1965 (morphology, varietal and mutant traits) ; CHAO, 1933 (principles and practice of Chinese rice breeding) ; COYAUD, 1950 (analysis of rice in Indo-China) ; GHOSE et al., 1960 (rice in India) ; GRIST,1965 (general monograph) ; Hu, 1961 (comparative karyology) ; IKENO, 1927 (rice genetics and symbols); ISO, 1954 (rice husbandry, 1936 (rice improvement in USA) ; especially in subtropic zones) ; JONES, JULIANO and ALDAMA,1937 (morphology of rice) ; KIKKAWA, 1912 (clas18 - Heredifas 55
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sification of cultivated rice) ; MATSUO, 1952 (genecology of cultivated rice), 1957 (Japanese rice cultivation) ; MIYAZAWA,1935 (mutations in rice) ; NAGAI, 1959 (japonica rice: breeding and cultivation) ; NAGAO, 1951 (gene analysis and linkage) ; NAKAYAMA, 1948 (mutations) ; PARTHASARATHY, 1938 a, b, 1939 (cytogenetics of Oryzeae and Phalarideae) ; PIACCO, 1936 (botanical classification) ; PORTERES, 1956 (agrobotany of cultivated rice); RAMIAH and RAO, 1953 (Indian rice breeding and genetics) ; ROSCHEVICZ, 1931 (taxonomy and agrobotany) ; V A N DER MEULEN, 1939 (natural crossing of rice in Java) ; VASCONCELLOS, 1963 (rice in Portugal) ; WORMER,1953 (photoperiod and morphology) ; YAMAGUCHI, 1926, 1927 a, b, 1931 (genetic analysis). Cf. also Studies on Rice Breeding in Jap. Journ. Breed. 4, 1954 (Supplement), by the Japanese Society of Breeding, Proceedings of the Symposium on Rice Genetics and Cytogenetics, Amsterdam 1964, sponsored by the International Rice Research Institute, Los Baiios, Philippines, lists of bibliography and other publications from this institute, as well as numerous articles in International Rice Commission Newsletter, Food and Agriculture Organization, Vol. I-XV, Regional Office for Asia and The Far East. (Note here the special issue 1963 of the Symposium on Rice Problems, held at the University of Hawaii, August 21-September 6, 1961.)
2. Taxonomy and phylogeny of cultivated rice The genus Oryza and related genera are probably derived from bamboo-like ancestors (CHANDRARATNA, 1964). The tribe Oryzeae was formerly placed in the subfamily Panicoideae but has now been transferred to the Pooideae. The tribe, consisting of circa 14 genera, is specially adapted to aquatic and swampy habitats. Some genera, among them Oryza, are considered to be of Old-world origin; others, like Zizania with the edible-grained species Zizania aquatica, are native to the Americas. Most genera have the basic chromosome number x=12, like the case is in Bambusa. Zizania, however, has the haploid number x=15, which might represent a secondary basic number ( c f . DARLINGTON and WYLIE, 1955). I n fact, the basic number x=12 of Oryza has also been considered to be secondary. In 1936 NANDI proposed 5 as the original number, from which x = 12 was derived by processes of secondary polyploidy. Other interpretations involve the derivation of 12 from x=7 (LAWRENCE, 1931) or a n origin depending on tetraploidy (CHAO, 1928) or
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Fig. 1. Idiogram of the mitotic chromosomes in Oryza satiua (by courtesy of dr. S. I<. SEN)
Fig. 2. Pachytene chromosomes of Oryza sativa (by courtesy of dr. S. K. SEN).
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amphiploidy after an hybridization of 5- and 7-chromosoined species with subsequent chromosome doubling (YAMAURA, 1933). The analyses by SHASTRY et al. (1960),as well as SEN (1963) and SHASTRY (1964) argue for a fully differentiated karyotype, consisting of 12 non-identical haploid chromosomes (Figs. 1-2). This indicates that the number 12 really functions as a sort of basic chromosome number, although the tribe Oryzeae itself may have originated from crossings between lowerchromosomed species and genera. Haploid individuals, with 2n= 12, show a n irregular meiosis, mostly with univalents, although one, two or more bivalents may occasionally be formed (MORINAGAand FUKUSHIMA, 1934; JONES and LONGLEY, 1941). Genetical evidence indicates a certain amount of gene reduplication within the genome (polyniery, homomery) , i.e. a more or less derived diploid constitution. In fact, HU (1964) by studying the mitotic chromosomes of a haploid plant reached the conclusion that 5 is a most likely basic number. He found a conspicuous similarity between some chromosomes of the karyotype, as well as a high degree of somatic pairing. There has been considerable argument about the number of species within the genus, depending on methods and means of classification. Recent workers seem to agree that there are circa 20 species; ROSCHEVICZ (1931) enumerated 19 species, CHATTERJEE (1948) 23 species. Both diploid (2n=24) and tetraploid (2n=48) species occur. Two species are cultivated: Oryza sativa L. and Oryza glaberrima Steud. The former, called "common rice", probably originated in SouthEast Asia (India and Indo-China, where there is a great diversity of variation). The latter, called "African rice", has its primary centre of variation in the Central Niger Delta, with secondary centres in other parts of Africa (PORTERES, 1956; KIHARA, 1959; CHANG, 1964; etc.). Archaeological findings date the antiquity of sativa rice in India back to 1000 B.C., in China to about 2800 B.C. Rice cultivation probably began ten thousand years ago. With regard to African rice the primary centre of Oryza glaberrima seems to have been formed around 1500 B.C. (PORTERES, 1.c.). Genome formulas like those in wheat and oats have also been worked out for the Oryra species. However, there is as yet no complete agreement. The following scheme is condensed out of reports by NEZU et al. (1960), TATEOKA (1963), CHANDRARATNA (1964), and CHANG (1964). The genome symbols are those recommended by The Symposium on Rice Genetics and Cytogenetics (1964, Appendix 2 ) , with some amendments by T. T. CHANG(in litt.) .

Species
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Genome symbol
Distribution
Diploid species:
I. 0. satiua L.
0. niuara Sharma et Shastry 0. rufipogon Griff.
0 . barthii Chev. 0. (perennis Moench subsp.) cubensis Ekman
AA
AA AA
AbAb AcuAcu
Cultivated; origin: Asia; now world spread Wild, annual; Asia Wild; widely intercrossing with 0. satiua; Asia Wild; Africa Wild; America Wild; Africa Cultivated; Africa Wild; Asia Wild: Australia
XI. 0. breuiligulata Chev. et Rochr.

(including 0. stapfii Roschev.) 0 . glaberrima Steud.
111. 0. officinalis Wall ct Watt
AgAg
AgAg
cc
EE
I\. 0. australiensis Domin

Tetraploid species
V. 0. minuta J. S. Presl et C. B. Presl 0. punctata Kotschy ex Steud.

VI. 0. latifolia Dew. 0. alta Swallen
BBCC BBCC CCDD CCDD
Wild; Asia Wild; Africa, Asia Wild; America Wild; America
Other wild diploid and tetraploid species are recognized. The discussion of them falls outside the scope of this essay.
3. Polymorphism within cultivated rice It is evident from data on pollen and seed fertility (NEZU et al., I.c., Table 2; O K A , 1964, Table 4) that there is a high, often almost complete fertility in hybrids within the satiua group (I), as well as within the glaberrima group (11), but that these two groups are greatly intersterile. We may, with some right, treat the satiua group as one species (ecospecies), and the glaberrirna group as another species (ecospecies). According to Hu (1960) the two groups have almost the same karyotype and show the same patterns of intragenomic pairing in haploid individuals. Much of the interspecific sterility seems to be caused by genic sterility, both in the haploid and diploid phase ( O K A , 1964). Univalent formation, the occurrence of anaphase bridges etc. indicate, however, structural differences, for instance chromosome inversions, as one cause of the failure of full bivalent formation. The situation is, however, more complex than just outlined, because each of the two groups is split up into subgroups also showing decreased pollen and seed fertility when crossed inter se. KATO and his co-
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workers (1930) were apparently the first to propose the division of Oryza sativa into an indica and a japonica subspecies. The indica type was represented by varieties from India, southern China, Taiwan, Ceylon, Java and other tropical regions, whereas the japonica varieties were found in Japan, Korea and northern China. Later on a special javanica type has been added, with affinities to both the preceding types. Although there is often a good deal of sterility in crossings between the types (subgroups), the division is losing in significance (CHANG, I . c . , ~ .24). The morphological and ecological features are not at all clear-cut. The sterilities depend to a great extent on the choice of the varietal combinations used in hybridization. Furthermore, the intense breeding work going on in all the countries concerned and the wide circulation of new varieties confuse the classification. We may reasonably conclude that different mechanisms bring about sterility: (1) purely environmental conditions or, rather, definite genotype-milieu interactions, (2) genes causing sterility, whether dominant or recessive, complimentary, duplicate or single, (3) structural differences, with deficiencies, duplications, small inversions and translocations, and (4) cytoplasm-genotype interactions. The classification of rice varieties was originally founded on morphological features, although sterility barriers were also considered. MATSUO (1952) made a thorough study of more than 1,400 varieties. He paid special emphasis to grain dimensions. Japanese lowland rice chiefly consisted of short-grained types, varieties from Sumatra, Malaysia, Borneo, Indo-China and India of long-grained types. Varieties with very long as well as broad grains were found in Japanese upland rice, in Java, North and West China, Europe and America. In his classification MATSUO combined the grain dimensions with 20 other morphological characters and could in this way distinguish three basic types of variation: One, the so-called A type, occurred to almost 100 per cent in Japanese varieties, one, the B type, represented to 90 per cent or more by Javanese varieties, and one, the C type, represented to 94 per cent by the Indian varieties. OKA(1953, 1954) in his scheme of classification introduced various physiological properties like resistance to salts, low temperatures and drought. A further physiological property, that of photoperiodism, has a considerable influence on population differentiation. BEST (1959) con(1964, sidered rice, in general, to be a short day plant. CHANDRARATNA p. 185 ff.) stressed the fact that rice varieties markedly differ in photoperiod sensitivity. Under natural day conditions the relatively insen-
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sitive varieties are, as a rule, early in heading and the sensitive varieties are late. However, this reaction also depends on what is called optimum photoperiod. CHANDRARATNA (l.c., p. 193, Table 57) showed the influence of three photoperiods (10.5, 11.5 and 12.5 hours) on various East Asiatic varieties when cultivated in Ceylon, Indonesia and Japan. KATAYAMA(1964) presented additional information on photoperiodism. Varieties which showed accelerated heading under short day conditions, were considered to be photoperiod sensitive and those, which did not show any acceleration, to be insensitive. Almost all representatives of the perennis group were sensitive. In cultivated rice such sensitive strains are considerably rarer. Of the two cultivated species, Oryza sativa contained more insensitive strains than Oryza glaberrima. The reason for the relative insensitivity of Oryza sativa may be its long history of cultivation. The geographical distribution of sensitive and insensitive strains was interpreted in terms of adaptation to natural conditions as well as to agronomic requirements. Thermosensitivity, in addition to photoperiod sensitivity, probably plays a great role in controlling varietal adaptability in rice.
4. Gene linkages and duplicate factors In 1959 NAGAO and TAKAHASHI recorded twelve linkage groups corresponding to the twelve haploid chromosomes of rice. Further data were then published by NAGAO et al. (1964). Well-known marker genes are for linkage group I: waxy or glutinous, wx, 11: liguleless, lg, and phenol reaction with the grain, Ph, 111: purple node, p n , IV: long empty glumes, g, V: purple stigma, Ps,VI: Daikoku dwarf, d,, and gold hull, g h , VII: undulate rachis, Ur, VIII: lazy growth habit, la, IX: supernumerary bracts below the panicle neck (Hokanuri neck leaf), nl, X: mottled leaf, bl, XI: brittle culm, bc, XII: glabrous glume and leaf, gl. Of special interest from a classification point of view is the phenol reaction gene in linkage group 11, since the reaction often distinguishes continental or Indian varieties from insular or Japanese ones. Some linkage groups are, in our opinion, not yet quite clear, since only one or a few genes have been identified with them. Duplicate genes (polymeric, homomeric, multiple genes) are often considered an indication of a complex genetic structure, for instance involving polyploidy (Triticum and Avena species, Arachis h ypogaea) or a derived basic number (Oryza) . However, also in a clearly diploid species like Pisum sativum (x= 7) polymeric genes, giving a segregation
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of 15: 1, have been found to occur. NILSSON-EHLE (1948) also considered the different erectoides genes in barley (x=7) to be polymeric. Segregation ratios used for a n evaluation concerning a n original or a derived diploid state must be regarded with much caution. Nevertheless, duplicate or even triplicate genes for one and the same distinct character have been described rather frequently in cultivated rice and may be taken to indicate a sort of hidden polyploidy or, alternatively, the occurrence of segment duplications, gene transpositions or within-gene duplications. Some examples in rice are: long versus short glumes, awned versus awnless, normal stature versus dwarfness, normal versus floating habit, shedding versus non-shedding grains, photoperiod sensitivity versus insensitivity, anthocyanin coloration of apiculus and leaf stalks, chlorophyll mutations of various kinds (albina, xanfha, virescent), resistance to Cercospora oryzae etc. In some cases, for instance the photoperiod insensitivity, the dominance relations appear rather complex. 5. Mode of reproduction Generally, cultivated rice is said to be autoganious. However, as reviewed by CHANDRARATNA (l.c., p. 61 ff.), there is a varying amount of spontaneous outcrossing. I n studies relating to conditions in the southern United States this was on a n average fairly low, ranging between almost nil and three per cent (BEACHELL et al., 1938). In Java VAN DER MEULEN (1939) studying indigenous varieties found natural crossing within variety groups to range between nil and three per cent, but between groups and in definite directions the percentage of outcrossing could reach values of almost 15 per cent. The same author also showed that the amount of outcrossing rapidly rose when the distance separating the parents decreased. I n the monograph of RAMIAHand RAO (1953) more data were presented. I n many instances of wild as well as cultivated rice outcrossing took place to a very high extent (in some cases up to 20 per cent or more), If the optimum humidity for development was reached without the requisite rise in temperature, spikelets niay still open although anthers do not dehisce. This observation has been utilized in artificial hybridization work . . . (l.c., p. 21). And further: It may be stated that external factors, weather conditions in particular, have a great influence on the extent of natural crossing as in Java, where up to 20 per cent incidence is reported. Certain varieties like the glutinous types, even very far removed, up to 30 feet, are known to contaminate varieties in
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and NARAYANAN the vicinity (p. 22). In a paper of 1963 SAHADEVAN NAMBOODIRIpresented low values of outcrossing in the winter season (around 0.1 %) but definitely higher values in autumn (0.6-1.2 %). It thus seems certain that spontaneous outcrossing is a common phenomenon, especially when different strains are cultivated close to one another. All caution is therefore required in exact studies on artificial mutation, either involving bagging or greenhouse isolation of the materials after mutagenic treatments or, alternatively, the analysis of specific marker genes, where contamination of materials and outcrossings do not prevent correct conclusions. Much work on artificial mutation does not fulfil these requirements in rice, as well as in oats and other cereals. Cultivated rice is an annual plant, although perennating habit, partially hereditary, occurs in some varieties (RICHHARIA, 1960 a). Vegetative propagation is fairly easily achieved by extra tiller production from dormant buds and subsequent separation of the tillers. Clones covering large areas can in fact be produced in such a way. In the case of utilization of haploids in mutation research this method may be very useful.
6. Aims of plant breeding Based on whether the rice crop is grown with artificial irrigation or with the help of precipitation only, the varieties of rice are divided into two categories, low-land (or wet) rice and upland (or dry) rice. Socalled floating rice is adapted to regions, for instance parts of Burma, Thailand, Vietnam, where flood water may rise more or less gradually as high as 20 feet. In such floating varieties the stems gradually extend with the rise in water level. So-called deep water paddy, an intermediate type, is grown in India and East Pakistan (cf. CHANG and BARDENAS, 1965, p. 23). Of the worlds three major cereals, wheat, rice and maize, rice has the highest productivity, although it is grown in a lesser area than that of wheat (PARTHASARATHY, 1963). According to recent FA0 statistics world production for the three cereal species expressed as means for the four years 1961162--1964165 is respectively: wheat-257 million tons, rice-253 million tons, maize-219 million tons. Rice is the chief staple food for about half of the world population. Far East Asia produces more than 90 per cent of all rice grown. Nevertheless there is an enormous range in productivity per hectare also in the Asiatic countries. Australia, Spain, Italy, Egypt and Japan rank very high in productivity,
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similarly certain parts of China. Thailand, India, Philippines, Cambodia and Laos, on the other hand, show average yield figures three to four times lower. The very low national averages can be attributed to a number of factors, for instance the use of low-yielding varieties, the planting of rice in unsuitable localities and regions, the lack of proper management of the crop. Without going into detail, it ought to be emphasized here that a more or less advanced plant breeding cannot be very effective without a proper development of cultivation methods and crop management. And, vice versa, improved soil and irrigation conditions, increased amounts of fertilizers are not effective without a n improvement of the variety characters relating to lodging resistance, utilization of soil nitrogen, disease resistance at high nitrogen levels etc. These improvements must thus go hand in hand. This is well illustrated by the contrast between indica and japoniccc varieties. The former ones are generally cultivated under fairly poor fertility conditions, whereas most present-day japonicas have been bred under conditions of high fertility. The two groups of varieties differ markedly in their capacity of absorbing and assimilating soil-applied nitrogen sources. Crossing work has been performed and seems to give promising results, although the cytogenetical differentiation of the groups renders a recombination of genes more difficult and protracted between than within the two groups. There are in the rice literature abundant mention of problems encountered in rice cultivation: poor soil, shortage of rain, flooding, lowyielding varieties, severe lodging, diseases, pests, insect damage of harvested and stored seed. All this makes further rice breeding necessary, in addition to practicing improved methods of cultivation. The many needed improvements require complex breeding schemes. On the other hand, because of the often primitive methods of cultivation, with their resulting low yield figures, a n improvement to a doubled or trebled yield per hectare should often be easily accessible. Questions relating to strain registration have also to be considered here. RICHHARIA et ril. (1960) inention that there exist thousands of rice varieties in India. In addition to the local varieties grown by farmers year after year, there exist in cultivation, according to the authors, around 450 improved varieties. New varieties are continuously being added. The existence of many improved varieties within a country inakes the organization of seed multiplication and elite production cumbersome. Moreover, the iniproved strains often deteriorate rapidly owing to intermixtures, outcrossings and mutations. The production,
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certification and distribution of improved seed is of immense importance, because without procedures of getting rid of low-yielding or otherwise inadequate local strains, yield improvement is impossible. The first step in plant breeding is a fairly simple one, viz. the selection of improved strains showing a wide adaptability to differences in climatic and edaphic conditions. Here problems of multiple cropping, of earliness, of photoperiodic insensitivity are in the foreground but, as mentioned before, also problems connected with soil fertility and irrigation. The aims of rice breeding cover numerous properties of the plant, morphological, physiological and ecological. They can be divided into the following headings:
A. Yielding ability This may be characterized as the output of high grain per day of cultivation. In the case of multiple cropping or to avoid the dry season improvement implies an accelerated output per day and season. Here the aspects of adaptation of a variety (race) to varying ecological conditions must be considered. Problems of adaptability, specific and general production capacity, and also of earliness enter the picture.
Plant type Several authors ( u . for instance BEACHELL and SCOTT, 1963; JENNINGS, 1964; TSUNODA, 1965; T. T. CHANG, 1965) emphasize the association of plant type and grain yield. CHANG (1.c.) describing experiments at the International Rice Research Institute concludes that, regardless of geographic origin, the nitrogen-responsive and high-yielding varieties have relatively short, narrow, erect and dark-green leaves. They also possess a stiff and short straw, fairly early maturity and medium tillering ability. Those plant characteristics permit light to penetrate into the region of the lower leaves which results in better light utilization, especially at high nitrogen levels. Most tropical varieties lack these traits. Here also the dwarfing or semi-dwarfing habit, appearing as a consequence of mutation or gene recombination, is to be considered. However, under adverse conditions in the tropics, or on dry land, such short or dwarf varieties are often less well adapted.
B.
Lodging resistance and strength of straw It is well-known from rice, barley, wheat and other cereals, that resistance to lodging is a complex phenomenon (RAMIAH and HAO, 1953; RAMIREZ and UMALI, 1956; CHANDRARATNA, 1964; T. T. CHANG, 1 9 6 4 ,
C.
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and others). In rice, lodging is referred to as root and stalk lodging or as breaking of the stalks at different times from the boot stage up to maturity or past-maturity. Furthermore, lodging resistance is correlated with root development and penetration, total plant height, length of the lowest and uppermost stem internodes, internode and node thickness (diameter), amount and lignification of sclerenchymatic tissues in different parts of the plant. The genetical back-ground is complex, but genes with major effects do exist.
D. Resistance t o flooding and saline conditions In this connection some features relating to rooting capacity and stem properties may be mentioned which influence resistance to flooding. There are varieties which even under complete submersion are capable of growth. In the case of stationary flooding water certain varieties grow and manage to reach above the water level. The rate of growth during flooding generally increases with plant age from two to four weeks; after the age of four weeks growth falls off, and in plants ten weeks of age the reduction in growth is pronounced (RAMIAH and RAO, 1.c.). To some extent the case of intermittent flooding differs from stationary flooding. In such a case the plants are subject to mechanical pressures of uprooting and washing off. This exerts a strain on stems and roots and requires a profound power of regeneration by the formation of secondary tillers when the main shoots have succumbed. RAMIAH and RAO (p. 200) conclude: The growing of a rice crop in the flooded areas is, under the present conditions, very precarious. In large areas of South East Asia rice is cultivated in land subject to salt water inundation or to a rise of the water table with subsequent formation of vast alkaline areas. Anatomical differences have been found between salt resistant and susceptible rice. There exists also a definite genetical background for salt water resistance. Salinity is especially injurious to young plants. When the seedlings have once established themselves, there is less trouble (RAMIAH and RAO, p. 204). The physiological aspect of saline resistance apparently lies in a high osmotic pressure of the cell sap, enabling the plant to withstand plasmolysis by the saline soil solution. ALIM et al. (1962) report the successful selection of a salt resistant variety on the shore of the Ray of Bengal.
E. Photoperiodism nnd ecrrliness These features are often described as physiological, with a genetically complex (polygenic) background. However, genes with major as well as
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niinor effects interact to give a certain end result. CHANDRARATNA (1.c.) considered photoperiodism in relation to earliness, us. lateness. He elucidated the fact, well-known to farmers in the tropical countries, that two types of rice exist: period-fixed forms, where the interval of seeding to heading is relatively unaffected by sowing-date, and date-fixed or season-fixed forms showing a pronounced effect of sowing-date (p. 183). Since numerous japonica-varieties are photoperiod- as well as thermo-sensitive, they cannot easily be moved southwards. Simple gene segregation is often noticed, with sensitivity as dominant feature, but conditions appear to be more complex than this (T. T. CHANG,in litt.). Probably many genes, and several alleles at each gene locus, are at work. Earliness in rice is to a great extent a consequence of the heading date. It may, according to CHANDRARATNA (1954) be partitioned into several components: (1) photoperiod sensitivity, (2) minimum flowering duration, (3) optimum photoperiod. The minimum flowering duration commences with germination and ends at a date determined by the genotype. Extreme values with regard to the initiation of inflorescences are 14 days, at one extreme, in the Japanese variety Eiko (BEST,1959) and, at the other extreme, 73 days in the Javanese variety Solo. Heading date in the photoperiod-sensitive varieties also depends on the time of sowing and the photoperiod of the season. Segregation for earliness is rather complex. Now and then major genes are found, recessive, semidominant or dominant. The number of gene loci with drastic effects on earliness is probably not low. In addition, however, there is a great number of genes with slight effects, just like in barley.
F. Quality Japonica varieties have generally a moist and sticky grain when cooked, whereas indica varieties cook dry and flaky. This also indicates the cooking preferences (or prejudices) in different parts of the world. BEACHELL and STANSEL (1963) have specially dealt with factors determining cooking quality. This is said to be connected with amylose content, temperature of gelatinization and setback viscosity. Colour, shape and size imply other preferences of the cooked product. Preference is usually given to white rice, although coloured varieties in some instances are higher yielding and nutritious, containing more protein and minerals than the corresponding white varieties. Rice protein is superior to wheat protein (RAMIAH and RAO, 1953), is more assimilable and richer in essential amino acids and in this respect comparable to animal protein.
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G . Disease and pest resistance The following pathogens, mostly fungi, cause severe damage in rice cultivation: Piricularia oryzae, causing blast, has almost universal distribution; Helminthosporium oryzae, common in East Asia, is a major disease in India and also occurs in the southern United States; Helminthosporium sigmoideum, occurring in Italy and USA; the conidial form of Sclerotium oryzae, causing stem rot, is virulent in Italy and USA; Cercospora oryzae, causing leaf spot, is common in America; Fusarium moniliforme, foot rot, common in parts of India; Ustilaginoidea uirens, false smut, common in the Bay of Bengal and Malabar. Bacterial leaf blight (Xanthomonas oryzae) is a destructive and common disease throughout tropical Asia. Hoja blanca, white leaf, is a virus with heavy damages on rice in some parts of the American continent. Virus diseases are increasing in form and prevalence (or are recognized as such; Ou, 1965). I n Japanese varieties resistance to the serious blast fungus is almost entirely lacking (YAMAsAKI and NIIZEKI, 1964), but outside Japan there are several cases of resistance reported. Early results of gene recombination involving blast resistance were described by IWATSUKI (1942) after hybridization of a Japanese paddy rice and a n upland rice from China. Numerous races of Piricularia oryzae are, known. This makes a continuous program of breeding necessary, until a gene for complete resistance has been found (if such a gene on the whole does exist or can be produced, for instance by artificial mutation). YAMASAKI and NIIZEKI (1.c.)studied the reaction of resistance to four blast races in 260 varieties of different species. The following survey is taken from their Table 3: Genetic constitution: T y p e of resistance to four races of Piricularia oryzae
4R
3R 1s
2R2S
1R3S
4s
Total
Oryza sativa Japanese varieties Foreign varieties Oryza glaberrima Other species
Total
1 96 1 2
4 32
13 18
6 12
54 21
78 179 1 2 260
100
36
31
18
75
In later experiments further materials of Oryza glaberrima and other species (with the genome formulas AA, CC, EE, BBCC, CCDD) were analysed. Non-Japanese varieties of Oryza sativa, as well as representa-
MUTATIONS AND CROP IMPROVEMENT. V I I
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tives of Oryza glaberrima, were frequently resistant. A program of recombination and back-crossing will no doubt be highly successful with regard to resistance, but much work is needed to combine it with high quality and productivity. On the contrary, a program involving artificial mutation may, carefully planned and handled, produce resistance more efficiently, especially in high-bred varieties. Several genes for resistance, dominant or recessive, single or duplicate, have been reported. Resistance to Helminthosporium oryzae behaves similarly. GANGULY and PADMANABHAN (1959) found in 538 analysed strains of cultivated rice, originating from India, China, Japan, Pakistan, Russia and USA, only six varieties resistant to this disease, four from India and two from China. The gene situation in this case is not as well known, but dominant simple genes for resistance, and probably also modifier genes, seem to exist. In general, resistance to fungus and virus diseases is of the dominant type, although recessive resistance is also met with. In this connection resistance to various pests should be briefly mentioned. PATHAK (1964) states that cultural and agricultural practices often offer possibilities for insect control and that spraying with insecticides gives immediate results. Biological methods of control, including that of plant breeding, have proved fruitful. Resistance has been found in several cases. Varietal differences in resistance (or rather various degrees of susceptibility) seem to exist with regard to the yellow stem borer (Schoenobius or Tryporyza incertula), the rice gall fly (Pachydiplosis oryzae) and possibly the rice eel worm (Tylenchus angustatus), whereas the data are less clear or even negative with regard to the serious attacks of the swarming caterpillar (Spodoptera mauritia), the rice hispa (Hispa armigera), the rice bug (Leptocorisa uaricornis), etc. Some important points relating to rice improvement have been touched upon in the preceding attempted analysis. There exist numerous other problems, of great significance, with regard to resistance to drought, low air temperatures at anthesis, dormancy of the seed at harvest time, factors of quality, like the incorporation of smooth hull, shattering of grain at maturity etc. In fact, improvement is probably more cumbersome in rice than in any other crop plant of similar importance, owing to the greatly varying edaphic conditions and the complex climatic situation of cultivation in the tropics and subtropics, the primitive and modern methods of management often found side by side,
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GUSTAFSSON -AND IVAR GADD

~ ~~~
the different requirements of quality (necessary demands as well as preferences and prejudices) and the enormous multitude of diseases, pests and other hazards. For an analysis of breeding methodology in rice, see CHANDRAMOHAN and PONNAIYA (1963). Gene recombination, accompanied by advanced methods of selection, testing and registration, will be the central technique in rice breeding for a long time to come. In recent years data have accumulated indicating that also methods of F, heterosis, polyploidy of the auto- or amphitype, and not least artificial mutation may be of help in future rice improvement. Genic and cytoplasmic male sterility has been detected (OKA,1964) and would form a basic condition for the heterosis method. Tetraploid rice strains have been produced by twin analysis o r colchicine treatment (MORINAGA, 1964). Sterility is often conspicuous in autotetraploid strains but its extent depends on the choice of parents, continued hybridization and selection ( O K A , 1954; MASHIMA and UCHIYAMADA, 1955). Amphiploid products involving species crosses, comprising Oryza sutiua, glaberrima, breuiligulata, latifoliu and minuta, have been analysed. So far, such amphiploid products lack practical significance but may ultimately lead to significant advances. The case of artificial mutation will be specially dealt with in the following chapters.
11. SPONTANEOUS MUTATIONS
Literature on mutations in rice is enormous. However, in most articles n o real analysis of mutation problems is performed. Numerous papers refer to isolated findings of mutant types. Often one is not sure of the correctness of experimentation or interpretation. Many so-called mutants may, in fact, be due to intermixtures or segregates after crossings. We d o not consider necessary, here or in the corresponding section on induced mutation types, to cite all publications dealing with mutations. The reader is referred to the literature list, where numerous papers not quoted in the text are entered. Spontaneous and induced mutations may be classified into the following groups: (1) Aneuploid chromosome aberrations (trisoniics, monosoniics, nullisomics etc.) . (2) Chromosomal alterations within the diploid karyotype: Deficiencies-losses of genes or gene sequences.
289
Ihplications-gains of genes or gene sequences. - Duplicationdeficiencies may arise as the consequence of meiotic crossingover or erroneous anaphase separation in translocation and inversion heterozygotes. Inversions-often difficult to detect, especially when small and paracentric. Translocations-commonly detected in the meiosis of heterozygotes (depending on the size of the interchanged segments). (3) Gene mutations-difficult to distinguish from small deficiencies (or duplications). The more refined the linkage map, the easier is the detection of deficiencies (two or more gene loci may be covered by one and the same deficiency). (4) Mutations outside the chromosomal genes, mostly in special organelles of the cytoplasm, for instance plastids, mitochondria, but possibly also in other plasmatic structures. Using viability of the mutated homozygotes as a criterion (since rice is chiefly a n autogamous plant), we may distinguish another series of mutation types, ranging from (1) lethal and semilethal mutants, represented by chlorophyll lethals (plastid mutations; WALLES, 1963), sterility mutants with complete or partial sterility (of the diplontic or haplontic type, MUNTZING, 1930), dwarfs of various kinds and vigour, (2) mutants with subnormal or normal viability, most of which are of no interest whatsoever from a breeding point of view, (3) high-productive mutants, possessing features of value for plant breeding (high-yielding ability, lodging and disease resistance, earliness, quality properties etc.) . Of methodological interest and indirectly connected with differences in productivity is the distinction between qualitative and quantitative mutations (morphological and physiological, macro- and micromutations, major and minor genes), although no sharp limit between the two contrary categories exists (cf. GUSTAFSSON,1965). With regard to changes in chromosome number (aneuploidy, haploidy, triploidy, tetraploidy) the reader is referred to monographs and articles by RAMIAH and RAO (1953), JONES and LONCLEY (1941), SAMPATH and KRISHNASWAMY (1948),JONES (1952),KATAYAMA (1963 a ) , MORINACA (1964). Many aneuploid types arise in hybrid materials, also in crosses between triploid and diploid individuals, as well as in the offspring of asyndetic homozygotes. Trisomic types are valuable in linkage studies. Chromosome-doubled haploids are particularly useful for the production of full homozygosity to attain purity of a strain. Similarly the haploids can be used with great advantage in mutation
19 - Heredifas 55
290
WKE GUSTAFSSON A N D IVAR GADD
research for the induction of soniatic mutations, specially in an organism like rice, since they can be propagated vegetatively. In fact, MELCHERS (1960) isolated mutants in haploid Antirrhinum which were not et al. (1942) have described obtainable in diploid materials. MORINAGA the origin of a diploid sterile dwarf mutant from haploid rice. and RAO summarized the cases I n their monograph of 1953 RAMIAH of spontaneous and induced mutations isolated up to that time. Chlorophyll mutations in rice are similar to the ones obtained in diploid barley (for references, see the literature cited, and for their classification, u. GUSTAFSSON 1940). Many spontaneous cases arise in hybrid offspring. Some segregate in a simple fashion (3:l ) , others in more complex ratios, for instance 15:1, caused by duplicate genes. The number of genes causing chloroplast breakdown is probably quite high. The phenotypes range from pure white (albina) uia yellowish (xantha) to light green or yellowish green (uiridis of different shades), striped types (striata, zebrina) , and colour changing types like lutescens, albescens and uirescens, spotted types (maculata) etc. CODD (1935) analysed a case of chlorophyll deficiency involving three recessive factors producing albina seedlings when homozygous. The variety Demerara Creole probably contributed two of these factors, the third one having been found in Mexican Edith, Americano 1600 and Blue Rose. Mutants displaying full or partial sterility are not rare. Interesting cases were described by NAGAI (1926) and MIYAZAWA(1835). NAGAI reported hereditary changes with pistils transformed into staminodia, with more than one ovary in spikelets of an ear, with parthenocarpy, with so-called paleaceous sterility, etc. Genes for inhibited bivalent formation (asyndesis) have been described (RAMANUJAM and PARTHASARATHY, 1935). Many cases of sterility have arisen in hybrid progenies. Dwarfness is often simple in inheritance and conditioned by a series of major genes (TAKAHASHI, 1964). I n general, the dwarfs are recessive (1923) ( c f . MATSUURA, 1933) describin character, although SUGIMOTO ed a case of a dominant dwarf. An interesting dwarf, arisen in a hybrid pedigree, was described by KHADILKAR(1960), the ear-head being compact and erect, with peculiarly suppressed internodes, with thick leaves, with lemmas shorter than the paleae. JONES (1952) described a mutant type with short internodes, intermediate (semidominant) in inheritance. Dwarfs were also reported by NAGAO (1951). A special group of semidwarfs (uiz. Taichung 1, I-geo-tze and Dee-geo-woo-gen) deserves special mention, as they have normal-sized grains and are highly nitro-
291
gen responsive and productive. The genetic analysis indicates the occurrence of one recessive factor as well as modifiers (T. T. CHANG,in litt.). Interesting mutants of the drastic qualitative type have been enumerated by many authors, for instance ageotropic mutants (JONESand ADAIR, 1938), anthocyanin mutants (RAMIAH, 1930, 1935), compactoid mutants with short stems and bushy growth (KADAM, 1932), a neckleaf mutant (JONES, 1952), a mutant type with long glumes (KADAM, et al., 1941). TERAO (1917, 1922) studied a partially sterile mutant with large grains. JONESin his paper of 1952 also referred to mutants showing a sort of discoloration, covering different parts of the vegetative (1929) system (leaves, culms, panicle branches, also hulls). ENOMOTO reported a mutation of the Lux-allele (waxy) to the corresponding+ allele (Wx). Types having coarse culms occur. Mutants without ligule, auricles and collar are on record. Mutants having rolled and twisted or narrow leaves, may be placed in the semilethal or subnormal group of viability. MIYAZAWA (1935, p. 109) described a case of extremely late ripening due to mutation: . . . no variety could be found which shows such a late season. Such mutants seem to arise now and then, independent of place and variety. A series of other traits, not studied in experimentally controlled mutant material, are referred to in the monographs and review papers quoted in paragraph I. 1 : General references; for instance NAGAO (1951) and CHANG(1964). Studies on induced chlorophyll mutations often include control data ( c f . p. 306). The sum of spontaneous rates of chlorophyll mutations reach figures of about 0.2 per cent, but occasionally higher figures (0.6 %) are reported. Data on reverse or back mutation were published by TERAO(1917, 1922). A recessive mutant strain with especially large grains gave when selfed a few normal plants (less than 10 %). Some fixed normal strains (homozygotes) and great many heterozygotes were found in the offspring of those normal plants. Mosaic types bearing large-grained and normal parts in a single plant were also found now and then. TERAO calculated that the frequency of the transformation from a to A in vegetative cells was about one per cent. In the generative tissue (germ plasm) the transformation coefficient was three per cent. The back mutation of wx (waxy) to Wx (normal) in pollen grains happened at a rate of 0.1 per cent (ENOMOTO, I . c . ) . According to the present writers, there is a great need for a careful analysis of spontaneous mutability in pure lines as well as in variety
292
AKE
and species crosses. If, for instance, the juponica and indica groups of varieties are really differentiated by a series of small structural alterations-and not only by gene mutations-there should be a high increase of mutation rates in the hybrid offspring, both involving the general genotype and that of specific genes. The order of mutability ought to pure lines. be: species hybrids > inter-group > intra-group hybrids Specific marker genes, like the wx-locus, or group genes like chlorophyll lethals, but also chromosome breaks leading to segmental interchanges, would form valuable test materials for such a comparison. It is probable that the increased mutation rates of inter-group and species hybrids will greatly contribute to the detection and isolation of useful mutations.
>
111. MUTAGENS AND THEIR ACTION IN THE FIRST GENERATION

1. Mutagenic agents
In their bibliography on effects of ionizing radiations, covering the period of 1896-1955, SPARROW et al. (1958) referred to 29 articles dealing with rice. Since 1955 the number of such articles has greatly increased. I n fact, CHANG (1965) argued that since 1957 more than 100 articles directly or indirectly related to radiation work have appeared in rice literature. During the same period about 20 articles dealing with the inheritance of quantitative agronomic traits in rice were published. The wide ratio between the two categories of publications clearly indicates that quantitative traits received much less attention in relation to their economic importance. The first studies on the use of ionizing radiations in rice cultivation go back to reports by YAMADA (1917), NAEAMURA (1918), KOMURO (1919, 1922, 1924, 1931, 1944) and SAEKI(1936), dealing with the socalled stimulating effects of X-rays on yield. KOMURO and SAEKIobserved that X-radiation may act as a positive stimulus at low and moderate doses but will damage at high doses. They also relate these effects to the moisture content of the seeds. Apparently ICHIJIMA (1934) was the first scientist to induce mutations in rice, applying X-rays and ultraviolet. His work was then followed by investigations of IMAI (1935 a, b) , KANNA (1935), IMAIand KASAHARA (1937), as well as by RAMIAH and PARTHASARATHY (1938). In Italy SAMPIETRO (1935) claimed to have induced mutations by ex-
293
posing plants at anthesis to electric light. After the introductory work carried out by Japanese and Indian scientists in the middle thirties a rapid development took place. The mutagenic treatments involve X-radiation on dormant or soaked seeds ; y-radiation given acutely or chronically to seeds and growing plants; neutron radiation on seeds, and, finally, treatments of seeds using radioactive solutions and different chemical compounds (ethylene oxide, ethyleneimine, ethyl methanesulfonate, etc.) . References to the kinds of treatments, doses, stage of development, first generation effects can be found, infer alia, in BEACHELL (1957), BEKENDAM (1961 a, b), BHATTet al. (1961), CAMPOS and VELASCO (1962), CHAO and CHAI (1961), FUJIIand MATSUMURA(1959), FUJII (1962), HORVAT (1961, 1962), HSIEH (1959, 1961 a, b), HUANG (1961), KATAYAMA (1963 b), KAWAI (1962, 1963 a, b), KAWAI and INOSHITA (1965), MABUCHI e f al. (1961), MARIE (1962, 1963), MASHIMAand KAWAI (1958), MATSUMURA (1965), MATSUMURA and MABUCHI (1961, 1964), MATSUO et al. (1958), MATSUOand ONOZAWA (1961), MATSUO and YAMAGUCHI (1962), MYTTENAERE ef al. (1965), NARAHARI and BORA (1963), NAYAR (1965), NICAISE (1959), OHTA e f a[. (1957), OSONE (1963), OUANG (1963), SUKANYA BAI e f d . (1957), SIMON (1963), SORIANO (1961, 1964), YAMAGATA and SYAKUDO (1960, 1963 b), YAMAGUCHI (1958 a, b), YAMAGUCHI and ANDO (1959), YEH and HENDERSON (1958, 1963). Most of the papers just quoted also contain data on mutability. In the following, selected papers will be discussed.
2. Radiosensitivity at the genotypic level
FUJII(1962) applied y-radiation from "'Co and ''Cs, as well as X-rays, to dry dormant seeds of 24 varieties of cultivated rice. Germination was scored two weeks after sowing. Doses applied were 20-50 kR. LD,, (for germination) amounted for Japanese and Chinese varieties to circa 40-50 kR, for varieties from Burma and India to more than 50 kR. FUJII also examined seedling height in various varieties one month after sowing and found LDso's of 40-50 kR in four Japanese and Taiwan varieties and definitely more than 50 kR in four varieties from Burma and India (Fig. 3). Parallel results to those of FUJIIwere reported by MATSUOef al. (1958). (Cf. below.) FUJIIalso presented data on other species of Oryza. The cultivated Oryza glaberrima is evidently rather radiation resistant (LD,, for ger-
294
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Burma 0
0
20
40 50
20
0
0
4050 Dose (kR)
20
i
Formosa 20 0
I
4050
Dose (kR)
Fig. 3. Differences in radiosensitivity of varieties in rice (redrawn after FUJII, 1962).
mination was above 50 kR). Wild (or semiwild) varieties and species on the diploid level showed corresponding LD,,s of more than 50 kR (0.sativa var. spontanea), between 20 and 40 kR (0.perennis, australiensis, subulafa) and below 20 kR (0.officinalis). Tetraploid species are rather radiosensitive, their LD5,s lying below 40 kR (0. latifolia var. alta, eichengeri) or even below 20 kR (0. latifolia var. minuta). To what extent the differences found depend on variations in seed size, seed shape, moisture content, difference in chromosome and et al., 1965) remains to be analysed. Accordnuclear volumes (SPARROW ing to FUJII the relative biological efficiency (RBE) of y - and X-radiations is approximately 1 :1.1.
MUTATIONS AND C R O P
IMPROVEMENT. V I I
295
9-
C .-
87-
e
0
CI
----
expected observed I
C 0
L rn
. c
654-
.o
Y
l Y
3-
10 20 Dose (kR)
30
40
Fig. 4. Differential radiosensitivity of segregates from a chlorophyll heterozygote in rice (redrawn after SHAMARAO and BORA,1961).
YAMAGUCHI and ANDO (1959)compared the radiosensitivity of diploid and autotetraploid rice (the Japanese upland variety Sensko). Before the y-irradiation ("Co) dormant seeds were stored in a desiccator over saturated sodium chlorate. Doses applied were 10, 20 and 30 kR. Dose rate was 1000 Rlhour. In general, the seeds of the autotetraploid suffered less than those of the diploid. This was especially clear with regard to seedling height (18 days after sowing), number of panicles per y1 plant at maturity and the fertility of the y1 panicles. SHAMA RAO and BORA (1961) found that X-irradiation of seeds, stabilized as to moisture content, of a n albina heterozygote (Aa) changed the segregation ratio from almost exactly 3 : 1 (P-value for agreement 0.99) to more than 9 : 1 at doses of 30-40 kR (P < 0.001). This means that the al6ina recessives (aa) suffer more from high doses than homozygous and heterozygous genotypes (AA and A a respectively). Two sets
296
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of experiments were carried out (Fig. 4), one comprising only segregating progenies (1), the other comprising progenies of heterozygous as well as homozygous genotypes (2). LD,,s for the genotypes were:
AA AAi-Aa aa
(1)
(2)
35-40 25-30
kR kR
10-50 30-40 20-30
kR kR kR
The differences possibly indicate an incomplete dominance of the genes for radioresistance, the order of resistance being AA > Aa aa.
>
3. Comparison between different types of ionizing radiations

BHATTet al. (1961) compared X and neutron effects on the segregation of the albina heterozygotes just described. The thermal neutron treatment (Brookhaven) ranged from 2.5X 10l2 to 2.5X lo*nth/cmz. None of the five neutron doses applied caused a 50 per cent decrease in survival. With regard to y-irradiation LD,, for the indica variety GEB 24 was 40 kH. Using X-irradiation it was above 30 kR and in the case of pile neutrons > 7.5X 10 np/cm2. MATSUOet al. (1958) presented data on X and thermal neutron treatments of dry dormant seeds representing four varieties. LD,,s for seedling height (16 days after sowing) and for survival at maturity were as follows:
Variety
I~Dfio Seedling height LDso Survival (at maturity) N
s
Hikuu 132 Norin 29
? 30 kR
-
Karalatli Pukhi
> 40 kR > 30 kR
> > >
3.6X108nth/~ni2 1.9 ,, 5.2 ,, 3.6 ,,
> 40 kR
-
2.4-3.GX1018nt~/cm2
> 1.9 >
40 kR 5 3 0 kR
3.6-5.2 3.6
,, ,,
,,
Seedling height showed less variation in the case of neutron treatment than after X-irradiation (Fig. 5), in agreement with results obtained by EHRENBERG and NYBOM (1954) for barley. Similarly, differences in radiosensitivity of soaked seeds are much less with neutron than with X-ray treatments. MATSUOet al. concluded that the decrease in fertility is linear for both types of radiation. Norin 29 is said to be more sensitive to thermal neutron irradiation than Rikuu 132. MATSUOand ONOZAWA (l961), using dormant seeds, delcl-iiiined the LD,, for seedling height and survival to cn. 30 kR of X-rays and
297
Rikuu No.132
No treatment Av. ht. 13,6cm
X - r a y s (LOkR) Av. ht. L.3cm
Thermal neutrons (5,2x1d3 nth/cm2) Av. ht. 3,6cm

0 1 2 3 L 5 6 7 8 9 10 11 12 13 1L 15 16 17cm
Karalat h
No treatment Av. ht. Il.Ocm
1 . ,
0 1 2 3 L 5 6 7 8 9 10 11 12 131L 15 16 17cm
X -rays (30 kR) Av. ht. 8.9cm
Thermal neutrons (3.6~10'~ nth/cm2) Av. ht. 9,2cm
Fig. 5. Frequency distribution of seedling heights after X and neutron irradiation. Note the smaller range of variation in the case of neutron as compared with Xirradiation. (Redrawn after MATSUO et nl., 1958).
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4 X 1018nth/cm2 of neutrons in the variety Norin 29. These experiments
were combined with chemical treatments using colchicine, diepoxybutane and 8-propiolactone. NARAHARI and BORA(1963) treated dry unhusked varieties of indica and japonica origin as well as hybrid selections with pile neutrons (Trombay). Four doses were applied: 0.1, 0.25, 0.50 and 1X 1013np/cm2. The highest dose was lethal, the two lowest doses showed little effect on germination and growth. LD,, seems to lie between 0.5 and 1.0X 10"nn,/ cm'. Certain spikelet abnormalities appeared in the first generation after treatment, mostly combined with sterility. OUANG (1963) compared the M, effects of y , X and thermal neutron treatments. Complete data are not presented. Four varieties were used. The LD,,'s for germination were: for X-rays 2 40 kR, for y-rays 50 kR and thermal neutrons 3.5X lO'*nth/cm'. Genotype differences in radiosensitivity possibly exist. SIMON(1963) working with Hungarian varieties found that dosages between 20 or 25 and 30 kR caused a 50 per cent reduction in survival and, simultaneously, a 50 per cent decrease in fertility. The author also compared the effects of y - and neutron treatments, applying doses of 25,000 R and rad respectively. Neutron irradi a t'ion was more severe than the corresponding y-treatment, especially with regard to root development. However, root destruction could be counteracted by the application of P-indolylacetic acid. Neutron treatment also produced a more profound MI sterility. YEH and HENDERSON (1963) compared the effects of y-rays from "'Co and fission neutrons (1.4X10'2 thermal neutrons +7.9X 10"' fast neutrons per cm2 and hour). Dormant seeds of the variety Nato (from a cross between japonica and indica rice) were irradiated. The results were the following:
<
<
LDjo for
Seedling height Survival Fertility
; ' 1956
y 1957 N 1956
> 40 kR > 50 kR > 4 hours

2-3
> 50 kR
-
N 1957
hours
2-3
hours
& 28 k R 30 kR 2 hours 1-2 hours
Some authors, especially KAWAI (1963 a and earlier), have applied solutions of radioactive isotopes ("P) to dormant or germinating seeds. KAWAI grew the seeds and seedlings in "P-solutions of variable concentrations for a period of two weeks (pH=5-6). The concentrations
299
ranged from 2.5 pGilseed to 40 pGilseed. The volume of solution was 0.6 cc per seed. After the treatment, the seedlings were washed in water, transplanted twice, given fertilizers and grown to maturity. One variety (Norin 8) was used. LD,, for survival lay between 10 and 15 pcilseed (as a n average for the years 1951 to 1953). Seedling height (after two weeks) and decrease in fertility gave similar LD,, figures. Root growth was more affected than seedling growth. After the atomic bomb explosion of Nagasaki in 1945 NISHIMURA (1957) gathered rice seeds from paddy fields situated 600-2000 metres from the center of explosion. In 1946 the next generation was grown for studies of morphological abnormalities and cases of decreased fertility. At the time of explosion the rice plants were approximately one month before flowering. Hence the grains gathered in 1945 ought to be compared with grains developing on growing irradiated y,-plants.
Distance from explosion centre Number of MI plants Number of abnormal plants
600 m 800 ,, 1,000 1,200 2,000 ,,

9,
7,
ca. 15,120 ,, 5,400 ,, 5,400 ,, 10,800 ,, 13,680
4.0 % 1.0 % 1.2 % 0.4 % 0.5 %
One thousand plants originating from the 600 metre material were carefully examined as to fertility. Sixty-one of them (6 %) had a n evident sterility. Most of the partially sterile plants produced partial sterility also in the next generation. Indeed, among 52 segregating families more than the half, or 29, showed a ring or chain of four chromosomes a t meiosis. Also trisomic and asyndetic individuals, as well as genetic dwarfs, were recognized in the offspring.
4. Developmental stage and effects of ionizing radiations The influence of moisture content and seed soaking on radiosensitivity was studied by MATSUO et 01. (1958) and MATSUO and YAMAGUCHI (1962). The results are on the whole in agreement with facts obtained EHRENBERC and others (u. the literature list). in barley by CALDECOTT, Soaked seeds were less sensitive to neutron than to X-ray treatments, relatively seen. Similarly, seeds with 6 per cent or 22 per cent moisture content showed a n increase in radiosensitivity as compared with seeds having 13 per cent moisture.
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401
OI.. ,
10 15 Seed water content

1965).
20
(lo>
25
Fig. 6. Fifty per cent decrease in seedling height after X-irradiation of rice seeds with varying moisture content. Calculated data (redrawn after MYTTENAERE et al.,
MYTTENAEREet ul. (1965, Table 1) irradiated rice seeds of different water contents (6.7-25.4 %) according to the equilibration method of EHRENBERG (1955; EHRENBERG and N Y B OM, 1954). X-ray dosages ranged between 5 and 50 kR. Five varieties were analysed showing conspicuous differences in radiosensitivity both with regard to germination rate and seedling height after 15 days. Striking contrasts were found with regard to the influence of water content. LD,,s for seedling height (Fig. 6) were calculated to be:
35 kR at 17 % seed moisture 3-4 kR at 7 % and 25 %.
A tenfold difference is thus found. Seed yield at XI maturity reflected this. The authors also studied the formation of free radicals by measuring net signal height at varying X-ray exposure and water content. At water contents between 7 and 11 per cent net signal height, i.e. values of irradiated seed minus values of control seed, was at its maximum and decreased at 17 per cent seed moisture. In his paper of 1965 MATSUMURAsuniiiiarized experiments with rcgard to the influence of the intensity factor on radiosensitivity. In one
30 1
series seeds were exposed to 5, 10, 20 and 30 kR of X-rays at dose rates of 10, 300 and 10,000 R/hour. Chronic, or rather long-time irradiation (10 R/hour), intensified the damage, for instance with regard to seedling height and panicle fertility. Storage of the seeds at a temperature of 15-19' C also increased the damage, both in the case of acute and chronic treatments. KAWAI (1962 a) irradiated growing plants at various stages of development using y-rays of eCo (2, 4, 8, 16 kR). In the 7,-generation the number of florets per panicle was markedly reduced when irradiation was carried out 25-30 days before heading, at a stage when panicle primordia are being formed. Fertility conspicuously decreased, most profoundly at high dosages, 10 to 20 days before heading, i.e. at the approximate time of meiosis. The same author (1963 b) also irradiated rice florets with acute X-ray doses of 0.5 to 4 kR 3-5 hours after anthesis, when the embryo is unicellular and the number of endosperm nuclei is 2 or more. The unicellular embryo stage was quite sensitive to irradiation as regards survival rate, although seed germination was only slightly affected. LD,, for survival lies at ca. 1.6 kR, which is a lower value than after acute y-irradiation at the heading stage. There also was a conspicuous difference with regard to the occurrence of partially sterile plants in the next generation. This is possibly due to a higher number of two-hit chromosome aberrations in the case of X-irradiation.
5. Chemical mutagens
So far only a few publications have dealt with the action of chemical mutagens in rice. The first paper available to us concerns the action of BAI et ccl., 1957) applied in powder form or acenaphtene (SUKANYA paste. Seeds were also exposed to a treatment with ceresan (a mercury seed disinfectant) in high doses (8-10 grams per kilogram seed), as well as to benzene vapour. The authors state that mutations arose and were tested for properties such as height, tillering, grain characters and yield performance. High-yielding types were isolated. The ceresan treatments are well worth repeating using controlled pure seed and all possible caution in avoiding intermixtures or crossings. In view of the often reported mutagenic effects of colchicine, especiaIly in Sorghum ( u . GUSTAFSSON, 1960, p. 15), the mutagenic action of acenaphtene has to be analysed more fully, not only in rice but also in other crop plants. MARIE (1962 a, b, 1963, 1965) induced mutations in some cultivars of
302
A K E GUSTAFSSON A N D IVAR G A D D
rice applying X- and y-radiations as well as chemical mutagens (diethyl sulfate, ethyl methanesulfonate), either separately or combined (0.5 % solution of both chemical compounds at 24' C). The treatment with ethyl methanesulfonate gave a survival after transplantation of 74 per cent of treated grains. Of this proportion 59 per cent were vigorous, 15 per cent were rather weak. A corresponding treatment with 48 kR (in two equal exposures with a rest period of 24 hours between the exposures) gave 33 per cent survival, with 28 per cent vigorous and 5 per cent weak XI plants. KAWAI and SATO(1965, unpubl.) compared the effects of two chemical mutagens (ethylene oxide and ethyleneimine) and X-irradiation. The material consisted of dry dormant seeds from a pure line, obtained after chromosome doubling of a haploid plant. X-ray exposure ranged from 10-30 kR. Ethylene oxide and ethyleneimine were given for a varying duration of time (EO: 2-8 hours, EI: 1-5 hours) in concentrations of EO: 0.1-0.6 %, EI: 0.02-0.5 %. The volume of solution was 25 cc for 100 seeds. After the treatment the seeds were washed and then, for germination, kept wet for 24 hours. Seedling height was measured 10 days after sowing in boxes in the greenhouse. Transplantation into the field was carried out one month after sowing. The fertility was determined from 100 spikes of 100 MI plants in each series. The results of the first generation were the following:
EO
EI
X-rays
LD,, for gerinina t'ion

LD,, for seedling height
LD,, for seedling height

LD,, for survival
50 % decrease in fertility
k0.5 % 2 hours 0.2-0.3 % 2 hours 0.1 % 7 hours 0.5 % 2 hours Not achieved
0 . 3 4 . 5% 2 hours 0.1-0.3 % 2 hours 0.04-0.07 % 5 hours k0.3 % 2 hours 0.1-0.3 % 2 hours
k25kR k 2 0 kR
20-30 10-20
kR
kR
MATSUO and ONOZAWA (1961) studied the effects of P-propiolactone, colchicine and diepoxybutane, alone or in combination with X-rays and thermal neutrons. Concentrations used were 0.25 per cent propiolactone for 60 minutes, 0.05 per 'cent colchicine for 24 hours and 0.1 per cent diepoxybutane for 60 minutes. The treatment with /?-propiolactone gave a 50 per cent decrease of germination and those with /?-propiolact-
303
one and diepoxybutane an approximate LD,, for survival at maturity. M, plant fertility was slightly but significantly decreased with colchicine and diepoxybutane, whereas a slight increase in fertility was noticed using 8-propiolactone. SHASTRY (1965, unpubl.) treated seeds of Oryza glaberrima with EMS-solution (concentration and duration of treatment not mentioned in the paper). 6. Translocations in the MI generation HSIEH (1961 a) made a comparison of quadrivalent formation in PMC's of five japonica varieties, seeds of which were treated with X-rays (20 and 25 kR) and thermal neutrons (3.6 and 4.9X 101anth/cmZ). With 20 kR the rate of PMC's having quadrivalents ranged between 29 and 45 per cent, with 25 kR between 32 and 50 per cent. The exposure necessary in order to induce one per cent of PMC's with quadrivalents was calculated to be 600 R. In the case of thermal neutrons, associations of four chromosomes occurred in 30-45 per cent of all PMC's applying the lower dose, and in 40-59 per cent at the higher dose. For the induction of one per cent PMC's with quadrivalents these values imply a dose of thermal neutrons of circa O.lXIO'anth/cmz. One R of X-rays would then in this case correspond to a value of 1.6X lO"nth/ cm'. (Such a comparison must be considered with all reserve.) There seemed to exist varietal differences with regard to the rate of quadrivalents induced. CHAO and CHAI (1961) irradiated equilibrated dry seeds of two varieties of rice (Taichung No. 65 and Chianan No. 8) with 20 and 25 kR of The X-rays and with thermal neutron doses of 2.7 and 4.0X 101anth/cmz. variety Taichung gave a consistently higher number of M, translocations than did Chianan. According to the authors 10 kR of X-rays were approximately equal in effect to 1.4X 1O1*nth/cmZ. This ratio closely corresponds to the one found by MATSUO et d . (1958). The induced chromosomal rearrangements resulted in a fairly high aniount of pollen abortion (between 26 and 66 % in Taichung and 33-53 % in Chianan).
IV. INDUCED MUTATIONS

1. Chlorophyll lethals as test mutations
Most scientists experimenting with induced mutations in Oryza sativa have commented on the origin of chlorophyll lethals: rates, types and
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proportions. In the following survey only special problenis relating to these lethals will be discussed. Following IMAI (1935 b) and KANNA (1935), IMAIand KASAHARA (1937) analysed 220 cases of X-ray induced chlorophyll mutants arisen in the offspring of 868 X, plants and 9,272 X, ears, i.e. an average mutation rate in panicle progenies of 2.3 per cent. The authors classified the mutations according to segregation ratios, either according to a n approximate 3: 1 proportion, with roughly 3/4 of all mutations, or to a more erratic and sporadic occurrence, with l / 4 of the mutations. Creamish and white phenotypes (albina sensu Into) comprised 56 per cent of the mutant cases, xantha six per cent and chlorina (uiridis) 17 per cent. Variegated and more rare types occurred to ca. 20 per cent. IMAI and KASAHARA further noticed the chimaeric structure of the X, plants, not all panicles of a n X, plant being heterozygous for a mutation. In fact, most cases involved only one single ear of a mutated XI plant. SHASTRY (1965) reported about chlorophyll lethals in Oryza glaberrima induced by EMS (albina, xantha, uiridis, tigrina numbering 34, 14, 12 and 1 respectively). Chlorophyll lethals were also recorded in a great number of wild species. Apparently differences occur in mutation spectrum (rates and proportions of the various mutant types). Tigrina, for instance, is generally the rarest type in Oryza brachyantha, where viridis types are common, but is the major type of chlorophyll mutation in "field spontanea (collection C-38)
'I.
A. Mutation rate and sequence of rice tillers Fig. 7 (OSONE, 1963) illustrates the development of primary and secondary tillers in Oryza satiua. The author studied the rates of albinn mutation in the various tiller progenies treating dry dormant seeds with y-rays from la7Cs(10 and 20 kR). The main tiller has the highest mutation rate. On the average, the mutation rate decreases with the order of (1) primary tillers, and (2) the order of secondary tillers. The secondary tillers give lower rates than the corresponding primary ones. Segregation ratios also change with the type of progeny. Primary tillers, in general, give lower segregation ratios than secondary tillers (10 kR: 7 7 0 albinas as against 18 %; 20 kR: 12 7 as against 23 70; in a fully normal segregation the ratio is 25 %). There is a clear difference, however, as regards primary tillers, too. The later formed tillers give segregation ratios which i n three cases of four were close to the normal 0.25 ratio.
305
Fig. 7. Development of primary and secondary tillers in rice (redrawn after OSONE, 1963).
Fig. 8. Size of meristematic tissues connected with primary and secondary tillering in rice (redrawn after OSONE, 1963).
OSONE (p. 4) concluded that the generative tissues of the later formed tillers are derived from a smaller number of meristematic embryo cells than the first ones formed. The smaller the number of partaking cells the larger the size of the mutated panicle sector and the better the segregation ratio in the second generation. The number of cells in the embryo initials forming the main stems is, according to OSONE,5-6 at most, and this number decreases with the age of the primary and secondary tillers. The size of the original cell groups of various tillers is schematically drawn in Fig. 8. B. Mutation rate in relation to dose intensity of radiation and seed storage MATSUMURAand MABUCHI (1964 b) , in their treatments with y- and X-irradiations of different intensities, found the following mutation rates using the sum of chlorophyll lethals as test mutations:
20 - Hereditas 55
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Dosc
DOSC
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ratc
GUSTAFSSON AND WAR GADD

No. of paniclc progenies
0
Rlutalioii rate\ Storage, hrq aflcr irr:idiatioii

250
.XI)
i.-,O
Chntrol 5 kR, S 10 kR, X 20 kR, X 6 kR, y 10 kR, y 20 kR, y
10 kR/hour
0.02 kR/hour
927 537 +657 842+777+657 777 $566 304+766+777+821 644 +606 393
0.64 7 0 ' 2.15 7 0 3.44 8.19 % 2.48 7 0 2.08 70 6.05 % 8.90 %
2.93 7. 4.50 % 6.54 % 5.06 7 0 2.70 70 2.55 70 5.44 %
' In comparison to the spontaneous mutation rate of pure lines in barley this is a high figure.
Mutation rates are for acute X-irradiation without seed storage circa 4X 10-8 per panicle progeny and R-unit. For chronic y-irradiation without seed storage it is circa 5X10-'. There is a certain intensification of mutability after storage, especially with the dose rate of 10 kR/hour at the medium exposure dose of 10 kR. The different types of chlorophyll lethals-dhina, uiridis, othersappear to 56, 26 and 18 per cent respectively. (BEKENDAMin his studies of 1961 b, Table 13, found the proportion5 to be 50, 30 and 20 per cent.) There is a slight difference in niutation spectrum with acute and chronic irradiation as well as after storage, but the reported differences, although partially significant from :i st:itistic:rl point of view, cannot be considered certain.
C. Stage of plant development and mutation rates KAWAI and INOSHITA (1965) compared the mutation rates after y treatments of various stages of development (I. tillering, 11. formation of ear primordia, 111. meiosis, IV. time of heading; cf. p. 301). In the 7,-generation only stages I to I11 gave chlorophyll lethals, with crlbina mutants predominating. The ?,-rates of I to I11 were, on the whole, proportional to the dose within each developmental group. Control and heading materials gave no chlorophyll mutations. Average mutation rates in the y,-generation were for the stages I to I11 4.2, 1.6 and 1 . 0 X respectively, calculated per R-unit and ?''-ear progeny . All four developmental stages produced chlorophyll mutations in the ?,-ear progenies (y,-strains). With some degree of reliability the values of mutability can be recalculated per ?,-ear progeny and compared to y,-ear progeny:
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Average mutation rates in the y,-genercltion recalculated per yl- and yp-progeny and K-unit
wprogeny ?,-progeny
Stage I Stage I1 Stage I11 Stage IV
1 6 . 3 1~0 - ~ 19.0 17.6 . 11.8
o m lo-
0.8 1.3 0.9
In the y,-generation the following mutation spectruni was obtained:

Albina Viridis Others Sum
of mutations
Number
of
yl-
ear progenies
Number of y3strains
I. Tillering 1I.Earprimordia 111. Meiosis IV. Heading
65 63 49 84
% % % %
25 24 27 11
% % % %
10 % 13 % 24 % 5%
94 122 76 56
99 129 80 80
2,739 3,026 1,080 1,089
Early stages of development (I and 11) when irradiated give between 60 and 65 per cent albinas. Irradiation at meiosis produces a smaller amount of albina, slightly more of viridis and considerably more of other types. Irradiation at the heading stage leads to a pronounced excess of albinas with few viridis and other types. The statistical analysis with regard to differences in mutation spectra gave the following results (following KAWAI and INOSHITA,Table 6) :
Tillering Ear Primordia Meiosis Heading
Tillering Ear primordia Meiosis Heading
0
%
0
%
*%*
It is evident from the data of y l - as well as y,-progenies that the dosemutation rates per rontgen are considerably higher at low dose exposures than at high ones, indicating a n elimination of mutated cells in all four stages of plant development, especially at the stage of meiosis. The authors conclude that the initials of egg and pollen cells of each spikelet are differentiated already at the stage of formation of ear primordia. They also state (p. 254) that a n irradiation at the stage of tillering, followed by a screening of yp-mutants, is a more effective method than raising also the ?,-generation of normal y,-plants. In this connection it ought not to be forgotten that the spectrum of chlorophyll
30X
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lethals clearly differs with the stage of irradiation, probably owing to processes of elimination of definite mutant cells. Although not yet analysed, this elimination may occur in the case of viable, respectively productive mutation types, too. In his paper of 1963 b KAWAIreported about mutation rates, applying a n acute X-irradiation of florets, 3-5 hours after anthesis before the first mitosis of the embryo. Exposure doses ranged between 0.5 and 4 kR. The spontaneous chlorophyll mutation rate in this case was 0.3 per cent. The average chlorophyll mutation rate per X, progeny and R-unit rose to 17X10-8 at 4 kR.
D. Mutation rates in the case of different mutagenic treatinents There are relatively few reliable experiments comparing different types of radiation. According to data published by MATSUO and ONOZAWA (1861) 30 kR of X-rays and 4X 1O1'nth/crn2, given to dormant seeds, induced 4 and 6 per cent of chlorophyll lethals respectively. CHAO and CHAI (1961) published data which give approximate figures relating to the relative X and neutron efficiency, with 10,000 R of X-rays corresponding to circa 1.4X 101anth/cmz, i.e. in accordance with data given by MATSUO and ONOZAWA. KAWAI and SATO (1965) compared the mutagenic effects of X-rays, ethylene oxide and ethyleneimine and found the maximal effects of ethyleneimine to be circa five times as high as the corresponding effects after X-irradiation. The maximal amount of mutations obtained with a treatment using ethylene oxide corresponded to a dose of circa 10 kR. With treatments causing 50 per cent reduction in plant survival the mutation rates were:
Ethyleneimine - 55 %, calculated per M, seedling - 10 70, X-irradiation ,, Ethyleneoxide - 4 %, >,
9, 9, 9 9, 9,
E. Chlorophyll mutations i n relation to other mutation types A careful analysis was carried out by KAWAI (1963 a) in treatments with solutions of radiophosphorus. In the M, generation he distinguished between four groups of mutations: (1) chlorophyll lethals, (2) cases of decreased fertility, (3) cases of morphological changes accompanied by sterility, and (4) morphological changes with full fertility. The mutation rates could be calculated in relation to the number of M, plant progenies ( a ) , M, ear progenies (b), M, seedlings (c) and mature M, plants (d). Cases of chlorophyll mutations can be related to methods a,
309
b and c only, whereas viable and productive types can be related to all four types of calculation (a-d). The increase in mutability after P-treatments was obvious, especially with regard to chlorophyll mutations. N o chlorophyll lethal was discovered in the control material. Concerning other mutant types in p. 12) stated: Frequency of total variated the controls KAWAI (lx., strains of control plots was also high and more than 20 per cent; however, in this case, the number of variants was very small and usually one variant segregated in individual variated strains. Many noted control mutants seemed to be modifications rather than genetic changes, as suggested by the M, and M, analyses, where the contrast between control plots and treatments was more pronounced (7 % mutated control strains against circa 100 % strains with genetic changes in the treatments). It is evident from the grouping of c + d (Table ti) that in the series of high dosages (15-20 ,uCi/seed) approximately every second mature M, plant is genetically changed. Probably the real figure lies higher. According to these data, chlorophyll mutations seem to be in the minority of the four groups of classified mutants. They comprise 4 per cent of all recorded instances if calculated on M, basis (Table 6) and 5-10 per cent on M, and M, basis (Table 14).
2. Sterility and dwarfness
Sterility and dwarfness are features reported by numerous scientists in mutation research. Often they are connected in such a way that genetic dwarfs are found to be more or less sterile, or, on the other hand, that sterile mutants are less vigorous, more stunted than normal. Induced sterility of the semi-sterile type may arise as a consequence of chromosome interchanges (see for instance PARTHASARATHY, 1938; OKA et al., 1952; NISHIMURA,1957; OUANG, 1958; CARPENAand RAMIREZ, 1960; SORIANO, 1961; HSIEH,1959, 1961 a, b; HSIEH ef a ! . ,1962). It may also appear as the result of irregular M, meiosis leading to the origin of, for example, trisomic or monosomic progeny plants (PARTHASARATHY, I.c., examining primary and derived trisomics; CHALAM et a ! . , 1959, examining monosomic dwarfs, following treatment with *P). CHAOet al. (1960) and KATAYAMA (1961, 1963 a and b) described sterile mutants, showing asynapsis (or desynapsis) at meiosis, with the origin of trisomics and other chromosome aberrants in later generations as a consequence of the partially inhibited meiotic pairing. In 1964 c KATAYAMA further described the occurrence of high sterility following a pro-
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ARE
nounced syncyte formation of PMCs. Sterile types of various appearance were also reported by ICHIJIMA(1934), SHAHet aZ. (1961), JONES (1952), YAMAGATA and SYAKUDO (1960) and KAWAI (1963 a, b) . In 1958 OUANG and CHANGclassified induced sterility into five different types with regard to degree of sterility and type of segregation but with no parallel cytological examination. Sterility may also appear as a consequence of rather common aberrations in flower structure ( c f . p. 290, NAGAI,1926), for instance distorted lemmas and paleae, stamens transformed into staminoids, changed stigmas, twisted spikelets, two ovaries (SHAH et al., 1961), etc. Similarly, aberrants with non-emerging panicles and cases of extreme tallness (JODON, 1958; SHAHet al., Z.C.; CHALAMet al., 1959) often show a more or less profound sterility of the diplontic type, using MUNTZINGS classification (1930). With regard to segmental interchanges the remark made by PARTHASARATHY (1938 a, p. 22) is of special interest: On a comparison with homozygous interchange races derived from experimentally evolved segmental interchange heterozygotes, as in Datura, maize and Pisum, where not a single case of position effect has been reported, it may not be different in rice. The back-ground for this statement was the occurrence of dwarf mutants in the offspring of semi-sterile interchange heterozygotes. In fact, HSIEH (1959) isolated 20 homozygous translocation lines out of 55 partially sterile heterozygotes displaying pollen abortion and decreased seed fertility. In 1961 he examined 14 translocation homozygotes, three of which covered entirely different chromosomes. Five of the 12 haploid chromosomes were involved in the homozygotes studied. HSIEHet al. (1962) studied the yield performance of 27 translocation homozygotes from three varieties. In one of the varieties plant height of the translocation lines showed a conspicuous increase in variance ( P = O . O l ) , possibly also in grain yield (P=0.05-0.01). Their analysis is of special value with regard to the further use of induced chromosome translocations in plant breeding. Genetic dwarfness varies in appearance from the most extreme stuntedness over semi-dwarfness to subnormality ( c f . p. 290). Grassy types and intensely tillering types have been repeatedly described. In general, dwarfness is accompanied by semilethality, although semi-dwarfness, et al., 1961) may be of value short-stature or short-culm plants (SHAH in developing non-lodging strains ( c f . p. 283). OUANG(1958) and OUANG and CHANG (1958) found that induced mutants affecting culm length most frequently were shorter than the parent strains used. In fact, in the variety Pei-ko, there were 1 7 cases
311
of shortened culm mutants against two cases with longer culms. The shortening, in general, comprised 10 to 20 per cent of the culm length of the parent strains. HSIEH (1962) isolated and studied numerous induced dwarfs in different varieties. Segregation was mostly monogenic, of the recessive type and regular. SHASTRY (1965) exemplified different kinds of dwarfs and short-stature plants. Especially interesting is a dwarf of Oryza sativa, resembling the wild species Oryza granulata. He also discussed dwarfness in relation to lodging resistance and productivity. With regard to the frequency of dwarf mutants in irradiation experiments KAWAI (1963 b, Table 17, p. 98) found that 11 per cent of the detected morphological mutants were dwarfish (culm length below a relative value of 70 %) ; 15 per cent possessed short culm (the culm length showed a relative value of 71-85 %), and 19 per cent of the mutants had somewhat shortened culm (a relative value between 86 and 97 %). Applying radiophosphorus in solution KAWAI (1963 a) found the corresponding values of 14, 9 and 27 per cent. In their earlier paper MASHIMAand KAWAI (1958) mentioned the occurrence of 3 per cent extreme dwarfs and 17 per cent dwarfs, similarly after P-treatments. In nearby fields of the atomic bomb explosion at Nagasaki NISHIMURA (1957) reported on the appearance of numerous dwarfs in progenies of seeds from paddy fields 600-2,000 metres from the centre of the explosion.
3. Viable mutants of morphological interest
Mutations without special plant breeding value but of morphological interest as gene markers have been induced in a considerable number of studies. Some are reported below. Ageotropisrn (RAMIAHand PARTHASARATHY, 1936) : An ageotropic mutant arose in X-ray experimentation. In the paper mentioned the mutation was reported as dominant but in their monograph of 1953 (p. 111) RAMIAHand RAOhave listed the mutation as recessive and due to a highly mutable gene. Ageotropic mutants are not rare in other plant genera as well: Zea mays, Hordeum, Pisum, Lupinus luteus and form a valuable material for physiological studies. Recessive mutants causing laziness were isolated by HSIEH (1962). Cf. also KAWAI (1963 a and b) with regard to approximate mutation rates. Narrow leaves (HSIEH,1962) : Such mutants are not rare in mutation work. They are often combined with dwarfness. The paper mentioned that a mutant with narrow and rolled leaves obtained after X-irradia-
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tion was found to segregate as a monogenic recessive. The so-called granulata-dwarf isolated by SHASTRY (1965) ought to be referred to here. Anthocyanin pigmentation (BASU and CHOUDHURY, 1961) : A mutant plant with a purple leaf-sheath was isolated in "S-experiments using a winter variety possessing a green leaf-sheath. The anthocyanin pigmentation was found also in stigma, lemma and palea, as well as in the awn tip o f the mutant. The pigmentation was said to be dominant in segregation and crossings. From a variety with purple apicule and stigma colour a "P aberrant arose showing loss of colour (CHALAM et al., 1962). Simultaneously awns were formed, and grain quality, colour and yield were improved. ( C f . NARAHARI and BORA (1963, p. 8 ) , concerning a neutron-induced early purple mutant). Black spot (JODON, 1958 a) : In a n X-ray experiment a mutant showing a black spot discoloration on the leaves was detected. The leaf spotting resembled attacks by fungi. The mutant homozygotes had fertile, well fitted panicles, although the effect of the spots on yield was not determined. This is similar to mutants found in barley and wheat and may possibly be due to changes in mineral requirements. Dense and sparse ears (KAWAI, 1963 a and b) : Such mutants are relatively rare in rice. In "'P-experiments they amounted to 0.2 and 1 per cent of the morphological changes isolated; after X-irradiation of florets the corresponding ratios were 0.4 and 0.4 per cent. A special category of dense-eared mutants consists of the so-called erectoides mutants (adopting a name from barley). This mutant type will be considered in the next paragraph. Short inner glumes (OKA, 1963): Possibly a recessive gene is responsible for this character, which varies in expression. Other cases of glume mutants have been found, for instance by YAMAGATA and SYAKUDO (1963). T a w n y l e m m a and palea (HSIEH,1962) : The lemmas and paleae are green at heading time but turn into a brown or tawny colour in the whole surface of the hulls at maturity. The mutant was induced by thermal neutron treatment and said to be conditioned by a dominant gene. Double kernels (MAJUMDAR, 1962) : In X-ray experimentation a n X, plant with abnormal vegetative and reproductive parts was isolated. The florets contained 1-6 additional structures between lemma and palea and had 1-7 ovaries, one or two of which gave rise to ripe small
313
kernels. The number of stamens varied from 4 to 12 instead of being fixed to 6. Long awns (SORIANO, 1961) : An awnless variety gave rise to mutant plants with awns more than twice the length of the grain (y-irradiation). Other cases, more or less questionable, are mentioned in the rice literature. Non-shedding (Hu et al., 1964): A non-shedding mutant obtained after X-irradiation was examined anatomically. The mutant lacked the abscission layer between the spikelet base and pedicel, in contrast to the parent strain which had a well-developed abscission layer. The nonshedding habit was said to be dominant, although the lack of a n abscission layer was recessive. The F, cross between mutant and parent is non-shedding and shows a n incomplete abscission layer. The mutant was dwarf-like, only 2/s of the normal plant height, and dwarfness was recessive. Non-shedding and dwarfness are conditioned either by pleiotropic effects of one and the same mutated gene or by changes in linked gene loci. Non-shattering mutant lines were discussed also by LI et al. (1962).
4. Productive mutations of plant breeding value
The earliest case found in literature concerning the direct utilization of mutations in agriculture dates back to the Chinese emperor KHANGHI who reigned from 1662 to 1723 (J. Heredity 1930, p. 282). Walking in a rice field he happened to notice a ripe, extremely early-maturing plant, with fine and full grain. The precocity of ripening was retained also in the following year. For 30 years it became the rice served on the emperors table. It was named Imperial rice and was the only kind that ripened north of the Great Wall. In the south, where the climate is mild and the soil fertile, two harvests a year could be obtained from it. It seems plausible to assume that the Imperial rice, or ya-mi, really arose by mutation. I n Appendix I of their monograph R A M I A H and RAO (1953) listed three spontaneous mutants which were considered to be improved varieties: Chanock sport (p. 316), Kitchili samba (p. 318), with the comments Fairly resistant to Piricularia; non-shedding and resists stem borer. A premier strain adapted to a variety of conditions and Senkuruvai (p. 322), said to be Fine early rice. With regard to induced mutation definite data on high-productive rice mutants released into practice are not available (u. the Addendum,
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p. 339). An early record of an induced valuable mutant is found in RAMIAHand RAO (Z.C., p. 248). They reported about 36 X-ray induced mutations affecting different characters. One mutant has proved useful from the economic point of view. It has a slightly shorter stature with a larger number of tillers than the original parent material, and has proved valuable in that it does well in rich soils where the problem of lodging is serious (quoting SRINIVASAN, 1941). MATSUO and YAMAGUCHI (1962, p. 246) listed four mutants under testing for local adaptation, three of which were induced by X-irradiation, one by P-treatment. One mutant strain (Saikai No. G4), equal in average yield to the parent variety, is said to be resistant to rice blast and stem rot. Another mutant strain (Fukei No. 54) has considerably higher yield than the parent (about 8 %), probably owing to an increase in panicle number, but is susceptible to blast. The mutant Fukei No. 53 has increased yield by 6 per cent, probably depending on a higher grain number per panicle, and is, in addition, characterized by a good stiffness of the straw. In a review paper KAWAI (1963 c) mentioned some practical results of mutation breeding, with regard to earliness, good grain quality, high yield, short and stiff culni, lodging resistance, high germinability under low temperatures, blast resistance. Some lines, as also mentioned by MATSUO and YAMAGUCHI (l.c.),are being tested on a large scale. Similar general remarks are found in a research note by JODON (1958 b) (also in Agr. Res., Wash., 1958, Vol. 7, p. 13) referring to an early mutant out of the Rexoro variety. It was characterized by an outstanding earliness in addition to the fine qualities of the original variety. It is unlikely that producers will ever grow this early mutant for the market, but other early varieties with the high quality of Resoro may be bred from crosses of the new strain with commercial rice. HEKENDAM (1961 a, p. 62) concluded that useful mutations were induced by X-rays and that it may be possible that several mutants will prove to be a direct improvement of the mother variety. The following useful mutant types were obtained: for resistance to Ccrcosportc, earliness, lodging resistance, seniidwnrfing, panicle length, size and shape of kernel. BOROUGHS (1962) reported about y-induced mutations affecting flowering date, length and rigidity of the culm, leaf width and grain length. I k m A and RAO (1958) similarly reported inutants with increased earliness and changed grain shape and size. CHALAMrt rtl. (1963) obtained two high-yielding mutant sclcctions in P-experiments, differing from
31 5
the parent material in panicle length, spikelet colour, awn properties, grain weight, apicule and stigma colour, besides having better grain quality. One selection (P 500-2-10-2-8) was superior to the parent by 13 per cent in 1960/61 and by 106 per cent (!) in 1961/62. SHAHet a / . (1961) selected 14 N,-lines obtained after thermal neutron treatment. One line (941), with normal panicles and grain, was short and sturdy and, according to the authors, might be valuable commercially or for breeding purposes. Another line (940-2) had a high tillering capacity. The authors also discussed hybridization of a short, stiff-strawed mutant (914), having coarse leaves but normal panicles and spikelets, with four different varieties. True-breeding, short-stature plants were recovered from all crosses in the normal ratio of 25 per cent. This characteristic may be of value in developing nonlodging varieties (p. 101). The short-stature mutations found in irradiated populations probably have practical application in the development of nonlodging rice varieties adapted to combine harvesting. Other mutant characteristics produced by irradiation that may have value in rice breeding programs are narrow and shorter leaf types and profuse tillering (p. 102). HU et al. (1960) X-irradiated ten varieties. Unfortunately, X, plants were raised without isolation. Some segregates from natural crossing were discarded altogether (p. 110). The material comprised circa 13,300 XI plants. In all, 269,000 X, plants were raised and 1,013 X, progenies were selected for further analysis. 148 out of 973 X, families were again selected. In two crops of 1958/59 18 promising lines were tested against the parent varieties. In the first crop season of 1959 two erectoides lines out-yielded the parent by about 20 per cent. Three highyielding selections surpassed the parent by circa 18 per cent. In the second crop season of 1958 one early-maturing line gave 15 per cent and 12 high-yielding lines on an average 12 per cent more than the parent. In the first and second crop of 1959 a special test of fertilizer response was also laid out, using three levels of N, P,O, and K,O, depending on the origin of the varieties. In the first season one erectoides mutant and one earliness selection were throughout inferior. One earliness and four large-panicled selections were superior, similarly 13 out of 16 highyielding selections. Some lines profited from high levels of fertilizers. In the second crop season two of five erectoides mutations and two of three multibranched mutations reached the parent. Four large-panicled mutants were inferior. One earliness mutant was equal in the first season and superior in the second season. Fifteen high-yielding selections were surprisingly good in productive capacity. Some high-yielding
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lines were superior in both seasons. A few indica mutants showed a decreased shattering. The chief result of the study was the finding that changes in ecological reaction could be induced, that the so-called erectoides mutants were sensitive to high levels of fertilizers and also sensitive to sheath rot (Acrocylindrium oryzae) , that selections in earliness and high productivity often showed a good yielding response in both seasons. Unfortunately the existence of outcrossings and intermixtures are not excluded. Further data on this series of experiments were published by LI et al. (1962). Here also yield data for the two seasons of 1960 were included. The authors stated: ( 1 ) Erectoides lines, so far obtained, are not specially valuable, either in jcrponica or in indica varieties. (2) An early line seemed to be rather promising. (3) Some high-yielding selections seemed to be promising, too. In fact, six such lines were chosen for further testing in the regional trials of Taiwan, all of them tolerant to high soil fertility and consistent in high yield in the crop seasons tested. (4) Large-panicled and multibranched lines were not specially good. (5) Tolerance for and utilization of high fertility could be induced. (6) Nonshattering lines were obtained. The authors also mentioned (p. 465) that X, lines resistant to rice blast were isolated. In several papers MARIE (1962 a, b, 1963) has studied the mutation method of rice improvement. In a series of mutagenic treatments some well-performing varieties, among them Allorio, a high-quality rice, were used (1957, 1958). 500 MI plants gave rise to 22,500 M, plants (1959), with 102 short plants selected. In M, (1960) out of 102 families 11 superior lines and 67 plants were chosen for continued study. From M, (1961) two families, with ten sublines and 88 plants, were carried over to the M, generation, where one family with eight sublines and 50 plants was again selected for the final trial. The best mutants were included in the ordinary recombination and transgression program. After y-irradiation numerous short-culmed, lodging-resistant mutants were isolated. Two mutants (Allorio Theta and Allorio Lambda) were high-yielding and lodging resistant; one of them (Allorio Theta) is slightly earlier, the other one (Lambda) definitely superior to the parent variety itself. A third mutant (Allorio Chi), induced by diethyl sulfate, was stiffer and higher productive than Allorio. In the M, and M, generations following P-1reatments MASHIMA and KAWAI (1958) isolated 283 mutant lines true-breeding and fertile. Of 69 lines tested in the next generation 15 lines were higher yielding than
317
the controls. Continuing the selection work nine lines were included in randomized yield trials. Lines with short culms, dense or sparse ears, late heading maintained a high yielding potential. After X- and y-irradiations OUANG(1964 a) selected 254 M, mutants, which after continued testing were reduced to 103 mutant lines in Ma, 39 mutant lines in M, and 22 mutant lines in M,. The mutants were then tested in regular yield trials (the second crop season of 1961 and the first crop season of 1962). Six lines were superior in the second crop of 1961 and six, partially identical ones, in the first crop season of 1962. Some mutants are characterized by vigorous growth and short stems. Insect resistance was found in one case. OUANGalso compared the results of segregation in irradiated and non-irradiated hybrid materials. According to him the variation in growing days, as well as in other characters, was larger in the case of irradiated than of non-irradiated reached the conclusion that for many practihybrid progenies. OUANG cally important properties irradiation leads to superior, i.e. quicker results, than hybridization alone. In a further paper OUANG(1964 b) indicated that of 254 M, mutants six cases were still under testing in the M, generation. Four mutants, with economic value, were specially studied and the yield per d a y was analysed in five generations:
Variety Earliness Yield per day Index of yield per day
Control (9CK) Mutant 110 77 123 ,, 195 Control (HCK) Mutant 207
102.2 days +2.4 ,, f 1 1 . 8 ,, +5.4 ,, 103.8 ,, f 7 . 4 ,,
41.08 kg 45.05 ,, 45.07 ,, 43.46 ,, 40.19 ,, 45.90 ,,
100 109.6 109.7 105.8
zoo
114.2
(+indicates increased earliness.)
Mutant 207, in particular, ought to be mentioned since it reached maturity seven days before its control and on a per-day basis produced considerably more, in fact six additional kilograms per day. It also produced a better total yield. In a further abstract (1964 c) OUANGconsidered that irradiated progenies can be bred true in 3-5 years, shortening by one half the time required for the conventional method of hybridization and back cross. SIMON (1963) described positive results using the Italian variety Maratelli. Other varieties, of Hungarian origin, did not give a similar positive response, either with regard to disease resistance or earliness.
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JJith doses of 24, 26 and 28 kR a conspicuous X, differentiation was obtained in Maratelli (the data are not absolutely clear with regard to the possibility of an original variety heterogeneity or the occurrence of intermixtures or outcrossings) . SIMONhimself concluded (p. 134) : Maratelli has proved to be a very mutable variety, although Italian authors could report mainly on negative mutations only. In fact, he was able to select remarkably early lines, one of which, for instance, ripened no less than 49 days before the original Maratelli variety. Under Hungarian conditions Maratelli did not ripen until mid-October, but the earliest X-ray lines as early as at the end of August. Special studies of induced curliness. - Induced earliness, or lateness of heading, are rather easily achieved (YAMAGATA, 1964). He found that variants for heading time were more frequent after y-irradiation of seeds than were chlorophyll mutants. Representative early and late heading y,-plants were studied with regard to earliness in the y,-generation. In his Table 1 (p. 33) the offspring of 21 y,-progenies were classified according to their distribution of heading date. Similar progenies have been grouped together in the following survey: TABLE 1 . Earliness to lateness, of 21 y,-progenies.
Earliness, days 2 4 - 1 6 8 Control
1247
4 -
8 346 ~(3)*-
159%
3progenies 4 >, 2 2 ,> 5 5

1, 11
18
53 41 19
19 115 32 16 8
26 52 18 15 102
38 17
8
1
11
I,
19 75 71 228
26 36 31 67 161
156 225 114 104
~~
Sum of progenies 21 18
113
190
II
213
77
31
1031
0 denotes the first heading date of the control. These three plants are considered more or less abnormal.
The analysis indicates that in the material presented 31 per cent of all y,-plants were earlier in heading than the parent (GimbBzu) and that 10 per cent were later. Moreover, it was relatively easy to obtain truebreeding early as well as late selections. a characteristic reEach variety used has, according to YAMAGATA, sponse to irradiation with regard to heading date. The varieties or
319
strains of early maturing type tend to produce more late maturing aberrants than those of late maturing type, and conversely those of late maturing type tend to produce more early maturing aberrants than those of early maturing type (p. 35). (The author erroneously writes late instead of early.) Thus, to a certain degree the mutation spectrum depends on the genetical constitution of the material used, a conclusion in agreement with recent results in other cereals, for instance barley. According to the author, the induced mutations for earliness analysed generally behave as recessives, but semidominance or even dominance may also be found. Mutations with major as well as minor effects are induced. Many early, or late, mutants are fully viable and productive. Heading date favourably lends itself to a profound analysis of the genetic back-ground of quantitative traits influencing earliness and yield. Yield and morphological deviation. - In his papers of 1963 (a and b) KAWAI dealt with productive mutants and their variability in various respects (for instance heading date, culm length, seed set density, 1,000 grain weight). In all, 283 true-breeding strains obtained in 8aP-experiments (1963 a) were analysed. After discarding mutant lines with abnormalities and greatly reduced yield 80 mutant types were compared with the control. Yield of these lines ranged from 64 to 111 per cent. Ten mutant lines had a yield index above 100 per cent and 29 mutant types above 98 per cent. The highest value was 111 per cent. With regard to heading date more mutants were late than early but some productive early types appeared. Number of ears per plant and florets per ear were often increased, similarly 1,000 grain weight. Culm and ear lengths, on the contrary, were usually decreased. In one mutant type 1,000 grain weight was increased by nearly 30 per cent. KAWAI (1963 a, Table 32, p. 45) listed 24 promising mutants. Two were very early (by circa 8 days) and gave, in spite of the earliness, relative yield figures of 97 and 99 per cent. The highest yielding strain (with 111 %) was similar to the parent in most characteristics but showed conspicuously increased tillering ability and was one day later in heading. After X-irradiation of florets KAWAI (1963 b) isolated 156 true-breeding mutant types. Several were considered useless from the practical point of view. However, 23 mutant types remained for analysis of i.a. heading date, grain properties, seed set density and yield. Early and late mutants were isolated although, on an average, less drastic than after *P-treatments. Twelve mutants with a relative yield index equal or
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Heading date
-8 (e a rly1
O -I
0 n
4 .
(late)
(No. of days)
No. of panicles per plant

80
100
130
Culm length
h -w
-w
. r 3
80
100
130
z
80
No. of spikelets per panicle
100
130
1000 grain weight
80
100
130
80
100
130
Fig. 9. Variation of six individual characters in mutant strains of rice selected for a productivity equal to or higher than that of the parent variety, Norin No. 8 (=loo). (By courtesy of T. KAWAI, 1966.)
32 1
above 100 per cent were listed (p. 101), five of which yielded up to 105 per cent or more compared to the parent. Short-culmed, high-yielding mutants were isolated. KAWAI calculated what he called deviation indices for 80 mutants from the P-treatment and 23 mutants from the X-ray series, with regard to differences from the mother strain in six characteristics (number of panicles per plant, culm length, panicle length, number of florets per panicle, seed set density and 1,000 grain weight). Slightly deviating types generally show high yields. Some early short-culm mutants, although rather extreme in deviation, were high-yielding. In general, in the group of distinctly different types, those with the lowest deviation indices, below an index of 40, gave fairly high yields. In his papers KAWAI also calculated the proportion of so-called beneficial mutants, i.e. such mutant types which are considered to be of direct value for agriculture and plant breeding. In the *P-treatments the average rate of beneficial types was 2 X lo-, i.e. two beneficial mutant types per tested 10,000 plants (1963 b, p. 66). With X-irradiation of florets the corresponding rate was twice as high or 4XlO-, i.e. four beneficial mutant types per tested 10,000 plants (1963 a, p. 115). The following data are of general interest: In the experiments using *Pa total of 478 mutant types were isolated. Beneficial types arose to 3 per cent (14 cases). Of them six lines (or 1.2 %) had a yield index of 103 or more. Similarly in the case of floret X-irradiation 287 mutant types were isolated. Of them 3.5 per cent (or 10 cases) were considered beneficial and 2 per cent (6 lines) had a yield index of 103 or more. We conclude, therefore, that approximately one or two of 100 mutant lines isolated and treated in the way outlined by KAWAImight be of value to agriculture and plant breeding.
5. Blast resistance and induced mutations According to a paper by Ou (1965) resistance to some of the major tropical diseases in rice, for instance blast (Piricularia oryzae) , bacterial leaf blight (Xanfhomonas oryzae) and some virus diseases (of the tungro type), could possibly be obtained by means of induced mutation. In many countries improved new varieties have been developed. Their performance is not always certain because of susceptibility to diseases. It would be of great value if resistance could be induced in these varieties. Little work has been done to explore these possibilities.
21 - IIeredifas 55
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1 t seems worthwhile to try, particularly in countries where rice research is more advanced and varietal resistance has been well explored. Scattered data can be found in the literature dealing with induced disease resistance in rice ( c f . this article, p. 314). However, a study by YAMASAEI(1966) ought specially to be considered in this connection. One variety (Norin No. 8) and one fungus strain (Ina 168) were used in experimentation. The rice strain is susceptible to all blast races so far identified in Japan. The fungus strain carries three or four nvirulence genes also identified in Japan. The rice materials tested were of the following sources:
(1) X-ray treatment o f seeds. The dosages varied from 5 to 30 kR. The progenies were ear-to-row and two-grain bulk progenies. The selection work was carried out in indoor growth chambers by the spraying method of inoculation with continued tests in further generations. In the first set of experiments six resistant strains out of 3,381 X, and X, progenies kept their resistance up to the X, generation, i.e. 3 number of two per 1,000 progenies. However, the blast resistance in these cases was not obtained from fully isolated X, materials. In a second similar experiment, where blast resistance was detected to a surprisingly high extent, outcrossing was not prevented. Consequently the data are not reliable.
(2) Gamma-irradiation o f seeds. Doses ranged from 10 to 80 kR. In this case the y,-plants were raised in an isolated paddy field in order to prevent pollen contamination. Inoculation was carried out in the field as well as in indoor growth chambers. The nest generation of screened seedlings was grown in paddy fields or in pots. Progenies o f surviving plants were again tested for blast resistance. Seedlings were rated for blast-resistance based on types and number of lesions. The frequency of yz strains segregating for chlorophyll mutations was about four per cent, for mutations with blast resistance 40 times less or 0.1 per cent. In general, growth chamber tests were more reliable than field nursery tests. Segregation ratios of blast-resistant plants in the y2 generation were low, since only single resistant plants (single strains in y s ) were found in the respective y , strains.
(3) Analysis of morphological mutants w i t h regard t o blast resistance. True-breeding mutant strains obtained after seed treatments using P, X-rays and pile neutrons and after proembryo treatment with X-rays were inoculated in field nursery and greenhouse tests. Of 888 mutant strains tested only three showed moderate resistance (0.3 %). Apparent-
323
ly mutations in morphological characters arise largely independently of mutations for blast resistance. (4) Gamma-irradiation of growing rice plants. Stages of plant development from post-meiosis to pre-fertilization were exposed to 10, 15 and 20 kR in a gamma-field. Self-fertilization was enforced by bagging. In later generations inoculation tests were carried out. In all more than 26,400 ya plants gave 44 screened plants. Only six of these were fully resistant in the next generation. This corresponds to a rate of 0.02 per cent. Owing to the kind of treatment (irradiation at late stages of plant development) very few recessive mutations can be expected in the yp generation. YAMASAEI considered the mutants found to be of the dominant type. Such mutants are necessarily much rarer than recessive ones. The chief previous result seems to be corroborated, though, that mutations for blast resistance can be raised using ionizing radiations, even when precautions are taken against outcrossing and contamination. Although frequency of blast-resistant mutations is low, utilization of these mutations in practical breeding should not be denied, because screening for blast resistance can be carried out on large irradiated materials at seedling stage. In several papers YAMASAKI and his co-workers (partially cited by YAMASAKI, 1966) have studied the mutability of the fungus itself. Using appropriate techniques they found that the pathogenicity can be changed both in a positive and a negative direction. Mutant strains with an altered pathogenicity can be used for differentiating rice varieties with regard to the genic constitution for blast resistance.
6. Biometry of induced mutations in quantitative traits (pol ygenes)

The expression polygenes is here used in the sense of NILSSON-EHLE (1913; cf. GUSTAFSSON and GADD, 1965), i.e. genes (factors, mutations) leading to small changes in the phenotype (micromutations), mostly in quantitative traits like height, kernel size, tillering, yield etc. The first analysis of induced mutations for quantitative traits in rice, based on biometrical analysis, was published by OKAet al. (1958). They chose for study a n easily defined quantitative character, readily selected for, viz. heading date, as well as another character, more diffuse and less easily selected for, uiz. plant height. Together with the control material two irradiated samples, given 6 and 12 kR, were propagated in
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bulks up to the X, generation. All plants showing recognizable changes were eliminated from the populations. Seventy-five X, lines were raised by random sampling from each population and laid out in a split-plot design with three replications. The authors concluded, after a biometrical analysis, that the populations did not differ significantly in mean values. Variability, however, was considerably increased in the irradiated materials. These two facts seemed to indicate that mutations with plus and minus effects have occurred in approximately equal frequency, giving a symmetrical increase of variability in both high and low direction. The environmental component of variance was much greater than the genetic one. If mutations in, say, 100 gene loci are responsible for the mutational changes, in both traits, then the mutation rate could be calculated to 2.4X lo- per r6ntgen for heading date and to 1.4X10- per rontgen for plant height. The increase in genetic variability was found to be 16 per cent per 1,000 rontgen for heading date and 14 per cent for plant height (Table 111). The conclusions of OKA et al. were criticized by BATEMAN (1959). After a thorough inspection of the material he formulated his opinion that there is no symmetrical induction of plus and minus mutations around the control means. In fact, with the dose of 12 kR heading date is significantly changed in a unidirectional mode towards increased lateness (P < 0.001). For plant height, both values of 6 and 12 kR border to the 5 per cent point of significance. It seems safest to conclude that there is a n excess of mutations in the positive direction (p. 427). Positive is used in the sense of increased lateness and plant height, which may be considered negative from a plant breeding point of view. In another experiment carried out by OKAet al. (1.c.) ten extreme X, lines were selected for both characters in all three populations, in high as well as low direction. Their progeny (in X,) were then tested by a split-plot design with three replications. Heritability values showed that selection for changes in plant height was effective in the irradiated samples but not in the control. With regard to heading date selection was effective in all three materials, although higher values were obtained for the X-ray populations. The existence of a selection effect in the control plots is possibly explained by a residual heterogeneity or (less likely) by recent spontaneous mutations. YAMAGATA in his paper of 1964, quoted previously (p. 318), made a further biometrical analysis of mutations of heading date. Doses of yirradiation of seeds ranged from 5-30 kR. y,-panicles were classified
325
.
:
i
t .
I I I I I I I I I I I I I I I I I
Heading d a t e ( A u g . )
Fig. 10. Frequency distribution of mean heading dates of individual X, lines after X-irradiation. I-IV relate to lines selected from different groups of XI fertility. V is the control. (Redrawn after YAMAGATA, 1964; each dot represents an X, line.)
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into four groups of fertility: below 30 per cent (I), 30-50 per cent (11), 50-75 per cent (111) and 75-100 per cent (IV) respectively. The populations were bulked separately from y2 to y1 with visible mutants discarded. I n the y,-generation, 60 plants were randomly selected from each of the irradiation and control groups and then classified with regard to heading date (Fig. l o ) , as well as culm length, panicle length and number of panicles per plant. Like OKA et al. (1.c.) YAMAGATA found no real difference in average heading dates between treatments and control. Wider ranges of variation were, however, observed in the treated groups as compared to the control. The author concluded that minor mutations were induced to both directions of earliness and lateness with similar degrees (p. 44). Moreover, YAMAGATA found that the genetic variances were positively correlated with the sterility of the y,-panicles and became generally highest in the most sterile group (I). Higher values of heritability were found with regard to heading date than for other characters, suggesting a rather pronounced effect when selecting for earliness (respectively lateness). Here it ought to be pointed out that in the material presented by YAMAGATA (Fig. 7, p. 43) 24 lines were earlier than the earliest heading line of the control (August 26th), i.e. 10 per cent, and 14 irradi a tion lines against one control line were later than August 29 (circa 5-6 %). The following table summarizes some of his data:
Genetic variance I11
(u2g1
Series
Control
IIeritability (h2) I11 Control
Heading date Culm length Panicle number Panicle length
0.981** 3.312** 0.065** 0.147
0.769 2.617** 0.022 0.033
0.210 0.545 0.061 0.036
0.401 0.206 0.115 0.177
0.381 0.161 0.041 0.004
0.156 0.057 0.104 0.075
GUCHI
Following a previous paper by YAMAGUCHI (1962 a) JALIL and YAMA(1964) analysed another quantitative character, uiz. grain size (length and breadth), after successive irradiations using y-rays, with and without selection. Seeds were irradiated with 10 and 30 kR. From each y,-plant four fertile ears were harvested. One seed was taken froin each ear, and all seeds of a population were bulked together and sown to raise a y,-generation. This was either left unirradinted or treated again with y-irradiation to give a y,/,-generation. A further irradiation was then carried out in the third generation with samples of the different populations. After single, double or three-time irradiations, the
327
populations were selfed, in this way giving rise to the following third and fourth generation materials:
Year
Control
10 kR
Y1
30 kR
Y1
1958 1959 1960 1961
Control
,,
39
99
Ya
Yl/Z
YZ
YliZ
Y1/2/3
Y2/3
Y1/2/3
Ya/o
Y3 Y4
y2/311
The series ys-10 kR, y,-30 kR and y,/,-lO kR were compared, on one hand, as well as the series y,-lO kR, yzl,l,-10 kR, y,-30 kR and y,/,1,-30 kR, on the other (examined series underlined). The y,-comparisons gave but little change of means as to length and breadth values. In the case of 11,-comparisons the length means were rather similar, but the breadth values decreased conspicuously after repeated irradiation (ynla/,- 10 kR and yp/,l,-30kR). "It is evident that the difference among treatments for length was not statistically significant while that for breadth was highly significant and that the mean value for breadth decreased with cycles of irradiation" (p. 152). I n another experiment selection was carried out for long and short grains, after successive irradiations. The selection was rather severe, 90 per cent of the y,-populations being discarded and only 10 per cent kept for continuation of the experiment. The material analysed biometrically in the y,-generation consisted of: Control-long grains Control-short grains
y,i,-lO kR-long
yzir-10 kR-short
grains grains
y21,-30kK-long
y,l,-30 kR-short
grains grains
Selection in the control material gave no effect. In the treated populations, on the other hand, the mean values shifted significantly in the direction selected for. It also is evident from the data that the mean squares for variance among plants for short grains were larger than their counterparts in the selection series for long grains:
Selection for Treatment Mean (mm) Mean square (among plants)
Long grain
7,
3,
9,
7,
Short gra i n
,,
3,
3,
Control y,/,-lO kR ;t2/,-30 kR Control ?v2/,-l0 kR 712/,-30kR
6.72 6.77** G.76** 6.72 6.60'#* 6.66""
0.058 0.083" 0.148** 0.081
0.237""
0.251**
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The authors drew the following conclusions: ( I ) ,4fter successive irradiation without selection the mean values gradually decreased, and the increase of variance was not specially high. (2) Selection for desirable variants would be difficult in this case. ( 3 ) Successive irradiation w i t h selection, on the other hand, shifted the mean values towards the desired direction and increased the variability significantly. (4) There were indications that the genotypic background plays a n important r81e in the manifestation of mutations in quantitative traits (cf. p. 319 of this paper). and S u z u ~ r(1964) reported about induced I n a recent paper SAKAI mutations for seven properties (plant height, heading date, number of tillers or panicles, panicle length, plant weight and grain yield). The material consisted of X,-lincs of a pure line (Norin 8 ) , seeds of which were treated with a rontgen dose of 20 kR, and was compared to lines of the control. In order to avoid effects of natural selection the two populations were propagated from the XI generation and onwards by the method of one plant-one seed offspring. Approximately 160 lines randomly selected from each of the two populations were planted in completely randomized blocks with two replications. Each line (i.e. row, plot) consisted of 1-1 plants, the two border plants being discarded at the measurements. Analysis of the genetic and environmental variances was performed. Corresponding covariances were obtained in order to study the pleiotropic effects of the mutations. Comparison between X, and control gave significant differences with regard to plant height, plant weight and grain yield but no significance with regard to number of tillers and panicles or panicle length. Plant height was significantly increased, but minus variants did occur. Plant weight and grain yield were significantly decreased, with a unidirectional minus variation. Heading date, number of tillers, number of panicles, panicle length were not shifted in any definite direction. In general, accordingly, it appears that X-irradiation effected niutation towards decrease in polygenic characters, except for plant height (p. 145). This conclusion overemphasizes the data presented. It is worth while noting in this connection that with regard to heading date no X,-strain was earlier than the earliest control line. This is somewhat puzzling since mutations in heading date are rather easily induced. The material analysed was not very large, however. I n addition, it may be pointed out that there was a wide variation in heading date also in the control material suggesting a possible original heterogeneity, although the authors themselves refer the wide variation in this case to spon-
329
taneous mutations (p. 145). C f . the contrary behaviour of the control material in the analysis by YAMAGATA(1964) using the pure line Aikoku. With regard to genetic variances, the authors concluded: ( 1 ) Also the control lines gave significant genetic variances (again an indication of original heterogeneity, outcrossing or impurities). (2) The genetic variances of X, lines were remarkably high compared to those of the control lines. (For some reason or other the genetic variances with regard to heading date are not presented.) (3) The environmental variances differed not much between X, and control groups. (They were large, relatively seen, in the cases of plant height, number of tillers and panicle length.) In their analysis of pleiotropic effects the authors attach much importance to the data indicating that the association of panicle length and number of panicles is the result of pleiotropic, but inversely operating effects of the genes. (There is in fact almost no difference in the size of the genetic correlations comparing X, lines and control lines, uiz, rc= -0.901 and -0.887 respectively.) It is well-known in cereal breeding that a negative physiological connection exists between amount of tillering and size of ear. This negative relationship cannot, however, in many species be considered absolute. This contrasts to the authors conclusion (p. 150) : As a matter of fact, these two characters as is known from experience, can be hardly combined in rice. They are . . . controlled by inversely operating pleiotropic polygenes, and accordingly, it may be almost impossible to improve further both components together. Certainly this is a difficult task, but by means of recombination or mutation the physiological contrast should be possible to break or at least to influence gradually. The authors further emphasized (p. 145) : It is also of interest to find that pc between number of tillers and number of panicles is not very high. It may mean that these two characters are to a certain extent controlled by different genes. The conclusion may be correct, but still the pG-value in this case is the next highest one (0.749). The relations between grain yield and four of its analysed components may be quoted here:
pc=0.849 for grain yield 0.530 ,, ,, ,, 0.469 ,, ,, ,, 0.085 ,, ,, ,,
and number of panicles (high) ,, plant height (medium high) ,, number of tillers (medium high) ,, panicle length (low)
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SAKAI and SUZUKI specially stressed the negative effects of induced mutations, also with regard to small phenotypic changes. The positive effects found with regard to heading date and plant height are interpreted as the results of minus mutations in vigour or growth of the plants. I n consequence the authors are critical to the conclusion reached by various authors ( c f . above) about a n increase of variability, of approximately similar size, in both the plus and minus directions. In our opinion they have overemphasized the general validity of their data, as well as disregarded the differences in behaviour found between different kinds of varieties (early uersus late, short uersus tall etc.). It is a fact that plus changes in heading date, to take one quantitative trait, do exist, combining increased earliness and yield ( N I L S S O N - E H L E , 1924, for spring wheat, after gene recombination, and GUSTAFSSON, 1963, as well as HAGBERG and PERSSON, 1964, for barley, after mutation and gene recombination). A few other mutation studies should be quoted here, since they were directly discussed by SAKAI and SUZUKI. In 1961 a MATSUO and O N O ZAWA published data concerning the induction of mutations after X-ray, neutron and diepoxybutane treatments. They found that the means with regard to culm length were almost identical after the treatments mentioned as compared to the control, although the variances were much increased, most in the case of ionizing radiations, less using diepoxybutane. Similar results were obtained with regard to grain weight. Generally, variability was increased in both directions. KAWAI et al. (1961) studied P-induced mutants, decreasing culm length. The mother variety, obtained after chromosome doubling of a haploid, was a pure line with long culms. I n the M,-generation approximately 4 per cent of the screened plants were short-culmed, ranging between 82 and almost 100 per cent of the control length. Sixty-eight strains were analysed for different properties and classified in plus and minus direction. Mutations increasing grain yield and earliness were obtained. The connection between yield and culm length is illustrated by the following survey:
Culm length Yield
Difference in yield significant (f)
X o . of lines
No difference from mother variety

3,
,,
>,
>,
3,
5,
,,
+
-
N o significant decrr:iw
9,
77
-t-
2,
1 2 3 9 6
33 1
120
0
@D
110@*
s
v
100em
e e e e m '0 e.n
e e e e e em
I ! aJ '5 90C .-
"S '8
0
m e
0 . 0
(3
80-
0 0 0 0
80 90 Culm Length ( O
100
M
Fig. 11. Connection between grain yield and culm length in 68 strains obtained in progenies of *T-treated rice. O=yield significantly increased, 0 =no significant difference in yield, 0=yield significantly decreased. (Redrawn after KAWAI et al., 1961.1
Significant decrease
9,
3, 3,
9,
9,
77
indicates increase in yield and SignifiCanCC of difference - indicates decrease in yield and no significance of difference
+ +
+
+
12 17 15 3
Of the total material analysed 26 lines ( = 3 8 %) showed the sanie yield level as the mother variety, or higher. Three lines (=4 %) gave a significant increase in yield, 15 lines ( = 2 2 %) a significant decrease. Three lines combined, as the survey shows, a significant decrease in culm length with a significant increase in grain yield. Obviously, in agreement with other experience of mutation viability, and in agreement with the data of SAKAI and SUZUEI, there is a surplus of changes in the negative direction. Remarkable is, nevertheless, the rather high
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incidence of short-culmed strains with an unchanged or even raised yield potential (Fig. 11). A conspicuous decrease in vigour and yield seems to be associated with culm length below a certain value. Finally, a paper by KAO et a!. (1960) may be briefly referred to. Four samples of rice seed were treated with doses of 16, 20, 25 and 30 kR of X-rays. The X, and X, data were analysed with regard to induced variability. No other selection was made except that X, plants showing sterility or extreme dwarfness were excluded. Heading date, plant height, panicle length, and some other characters, were considered. According to the authors, the data indicated that in the X,-generation the variances increased parallel to dose. In the X,-generation, however, the largest value was obtained with 20 kR and the variance then decreased again with 30 kR. The authors planned to continue their studies of induced variability.
7. Screening methods for the extraction of productive mutations
I n a paper of 1962 YOSHIDAanalysed the statistical background of different screening methods for mutant detection in rice and barley. His idea was to establish methods by which desirable mutants could be isolated with a minimum total of progeny plants and, consequently, a minimum amount of area, labour and cost. He compared the number of M, and M, plants necessary using the following five models of analysis: A-the conventional method applied for instance by NYBOM (and GUSTAFSSON) in Swedish barley mutation work, i.e. the entire number of kernels per RI, plant (or ear) are sown in a row (maximizing the number in barley to 20 kernels per row). B-the improved ear-to-row method. The probability of finding one M, line with at least one desirable mutant is denoted as P,. In similar way P, denotes the probability of detecting at least one desirable plant among n plants in a n M, line with one or more desirable mutants. The method in calculating in, n and nin when P,=P, implies, according to YOSHIDA, the improved ear-to-row method. C-one-plant-one-grain method, i.e. one kernel is taken from each MI plant (or ear) and all the kernels sown as one progeny. In methods and E two and three kernels respectively are selected ironi each M, plant (or ear). The conclusion of the author is that each of the new iiietliods 13, C, D and E :ire applicable in radiation breeding. Method A should not
333
be used in any case (p. 95). When discrimination of a mutant is difficult, method B will be useful. When a mutant is easily distinguishable, method C, with the MI plants grown densely, will be superior. In some instances of radiation breeding, according to YOSHIDA, methods D and E are preferable to method C. With regard to the number of M, plants (m), total M, plants (mn) and total MI and M, plants in combination (m+mn), required to obtain at least one desirable mutant, the statistical calculations indicated the following order of plants needed: M,plants (ni) C > D > 4 > E > 3 M,plants (mn) A>B>E>D>C MI and M, plants (m+mn) - A > B > C > D > E
YOSHIDA(l.c.,p. 110) also presented data with regard to the number of plants necessary in order to breed a new variety. According to him, using the conventional ear-to-row method, a hopeful strain of rice (Saikai No. 64) required for its isolation a total population of 45,284 plants and a total area of 2,446 m. If methods C, D or E with a dense planting in XI, or if method B is used, radiation breeding will probably, YOSHIDAconcludes, require a smaller total number of plants and a smaller total field area than required in cross breeding, although this depends to some extent on the treatment in later generations. (In our opinion such a conclusion is meaningless if the aims of improvement are not stated.) In fact, what YOSHIDAdenotes as new methods, was already proand LEIN (1943, p. 276ff.). They compared posed by FREISLEBEN method A (eine moglichst restlose Erfassung aller Mutationen) with mehod C (Auslese auf der Grundlage einer moglichst grossen Zahl von Nachkommenschaften) . They arrived at the conclusion, statistically and empirically founded, that the latter method was superior to the former. In his paper of 1951 GUSTAFSSON pointed out that there are certain short-comings with the methods C (D and E) in order to effect a n accumulation of mutations after repeated irradiations. The draw-back of the einkorn method is that there easily occurs a negative selection for poorly tillered plants, when proceeding along the plant basis, since plants with one tiller are represented in the same way as plants with several or numerous tillers. A selection on the spike basis may cause a negative selection, too, since here sterile and fertile spikes are similarly represented (p. 278).
334
ARE
In p r o g r a m of improvement by means of induced mutation there ought to be a careful evaluation of desired properties already before the mutagenic treatment. It is of no practical interest, whatsoever, to try to cover the entire mutant spectrum obtainable, including lethals and semilethals, in autogamous species. In order to isolate a recessive or semirecessive mutant of a special desirable type we need the mutated homozygote itself (or the corresponding heterozygotes) A segregating progeny of 10-20 plants, in the case of a chimaerical MI structure, is required in order to find a recessive mutation with some certainty. Since in numerous instances parallel deleterious mutations are induced, mostly in different chromosomes, the breeder must separate the desirable mutation as much as possible from deleterious back-ground mutations. For this reason, the ear-to-row method combined with an adequate number of M, progenies and M, plants per progeny, is definitely superior to the one-plant-one-grain method. In 1951, discussing repeated irradiation, GUSTAFSSON proposed a n alternative procedure. A certain seed material is irradiated, machine sown and, in the case of self-fertilizing species, bred as a population for a couple of years. Mutations are forced to accumulate by irradiation of population samples year after year. Finally the population is broken up and analysed with respect to its heterogeneity. With this procedure the unintentional selection for delerious mutations is counteracted to some extent. The method can be modified in various ways according to the population structure of the material and the aim of the project. In their paper of 1956157 GUSTAFSSON and WETTSTEIN stated for barley that with a seed irradiation of 10,000 R probably every plant raised from a n irradiated seed would give rise to one or more mutants in its progeny. The dose of 10,000 R in barley corresponds in rice to approximately 30,000-40,000 R. It is here postulated that spike (or panicle) progenies treated with a n adequate X-ray dose (or corresponding doses of other mutagens) give rise to 100 per cent mutations and that the segregation ratio of a mutant is not 3 : 1 but 4: 1 owing to a chimaeric structure (in fact, it frequently is 5: 1 or less). Then, on an average, at least 5 X, plants are required in order to obtain a recessive homozygote. Using the denotions of YOSHIDA(p. 96) and the formula advanced by FREISLEBEN and LEIN (p. 277), the following mutation frequencies for one-grain (C) , two-grain (D), three-grain (E), five-grain (F), ten-grain (B) and 20-grain (A) progenies can be calculated:

No. of x, progenies (m)
K O , of x, individuals (mn)
335
No. of
I.ecessive
Method
No. of X, and No. of grains X , plants per X, progeny (m+mn) (n)
XI
mutations
C D E
17
B A
c
D E F B A
10,000 10,000 10,000 10,000 10,000 10,000 100,000 66,667 50,000 33,333 18,182 9,524
10,000 20,000 30,000 50,000 100,000 200,000 100,000 133,333 150,000 166,667 181,818 190,476
20,000 30,000 40,000 60,000 110,000 210,000 200,000 200,000 200,000 200,000 200,000 200,000
1 2 3 5 10 20 1 2 3 5 10 20
2,000 3,600 4,900 6,700 8,900 9,900 20,000 24,000 24,500 22,333 16,182 9,429
With regard to the number of X, plants (m) needed to induce similar rates of recessive mutations in the different methods the order is C >D E > F > B A; with regard to the number of X, plants needed (mn) it is A B F E D C, and for the XI and X, aggregate (m+mn) it is a A B C > F D E. We may consequently assume that around three kernels per spike progeny (model E) would be the least expensive method in radiation breeding. There is the fact, however, that in the case of models B and A approximately two and four mutated individuals are obtained per mutation case. In avoiding a onesided selection for negative mutations, or freeing a desirable mutation as much as possible from harmful back-ground mutations, methods A and B seem the safest ones for plant-breeding purposes. Furthermore, the possibility of obtaining heterozygotes of a certain positive X, mutation is greatest in methods A and B, intermediate in method F, less in method D and E and nil in method C. Needless to say, the einkorn method may turn out to be very helpful when the breeder has no special aim of plant improvement for his experimentation or when he wants to study line differentiation with regard to small changes in the phenotype, which cannot easily be detected in segregating progenies by the naked eye. The preference of a special method cannot be determined once and for all. The decision rests on the plant breeder, his program, his resources.
>
> > > > > > > > > >
336
AKE
SUMMARY AND CONCLUSIONS Cultivated rice comprises two species: Orgza sativa (or common rice) and Oryza glaberrima (or African rice). Both are diploid with 2n=24. Hybrids between them are sterile. The situation is, however, more complex than so, since also 0. sativa is divided into groups of biotypes more or less intersterile. In spite of the sterility barriers the subgroups of 0. sativa are only considered to be subspecies by most geneticists. Two such subgroups consist of japonica and indica rice, after their chief centres of distribution. In addition, a javanica type has been proposed. According to recent experience this division of 0. sativa into distinct subspecies is losing in significance. Much discussion has centered around the basic number of rice, whether the number x=12 is original or derived. Some authors consider 5 , or 7, to be the original basic number, from which 12 is derived by processes of secondary polyploidy. Other authors consider that x=12 is the result of amphiploidy involving crosses between 5- and 7-chromosomed parents, or that 12 is a derived tetraploid number. Numerous cases of duplicate genes certainly indicate a kind of derived condition, but, on the other hand, there is little doubt that the two chromosome sets of rice have become greatly diploidized in the history of the genus. Cultivated rice is generally considered to be autogamous. There are abundant data available, however, indicating that under certain exterior conditions outcrossing may appear to a rather high extent and that it varies with the genetical constitution. In fact, in certain environments such outcrossing may rise to 15 or 20 per cent. The common rate of outcrossing is much lower, often less than one per cent. In careful experimentation on induced mutation all caution is necessary in order to avoid not only intermixtures, but also outcrossings and segregates. This is especially true with regard to the first and second generations after irradiation. It is a regretful fact, illustrated in the preceding chapters, that also prominent scientists have failed in this respect. When working out rules and methods for the induction and isolation of high-productive mutants it is advisable to use a genotypically pure line, preferably containing specific marker genes, and then to keep it isolated during experimentation and to work as cleanly as ever possible. Spontaneous mutations are not rare in cultivated rice, although careful analyses are scarce. The present authors consider it worth while to study spontaneous mutations and their frequencies in pure lines compared to variety and species crossings. Relying on information in other
337
plant species, for instance Triticum,Pisum and Antirrhinum, we may possibly conclude that mutability is at its highest in species hybrids, decreases in inter-group and intra-group hybrids and is lowest in pure lines. Chlorophyll mutations are specially suitable for such studies. Also individual gene loci ought to be studied more closely. A valuable marker gene in mutation research is the waxy or glutinous gene (WX) of linkage group 1. The induction of mutations in rice by means of ionizing radiation PARTHASAgoes back to work by Japanese and Indian scientists (IMAI, RATHY, and co-workers). In late years y , X and neutron treatments of seeds have been applied and compared, but also acute and chronic treatments of growing plants under various stages of development. Radioactive isotopes, like P, have also been applied. The dose giving 50 per cent lethality, seedling height and sterility varies with the variety, the water content of the seeds, the type of seed storage, the kind of atmosphere during treatment etc. For instance, in seeds with 17 per cent moisture a n LD,, of 35 kR for seedling height was found (MYTTEN A E H E et a)., 1965). In seeds with 7 or 25 per cent moisture it is less than 5 kR. Chemical mutagenesis has been specially studied by KAWAI and his co-workers (cf. the literature list), involving the use of ethylene oxide and ethyleneimine in comparison with X-irradiation. Other experiments on chemical mutagens include diethyl sulfate, ethyl methanesulfonate etc. Much work along this line still remains to be performed. Mutations induced in rice generally belong to the following types : (1) Chlorophyll mutations, (2) mutations causing sterility, semilethality and dwarfing; here also segmental interchanges, giving rise to partial sterility when heterozygous, but being highly productive when homozygous, ought to be mentioned, (3) mutations of the morphological kind, varying in appearance, most of which are useless from a practical point of view but often interesting with regard to the phylogeny of the species and its mode of differentiation, (4) productive mutations of definite plant breeding value, with regard to earliness, lodging resistance, increased yielding ability, changed photoperiodism, plant type, kernel shape, semidwarfing etc., (5) mutations with increased resistance to diseases, for instance blast (Piricularia oryzae), and (6) productive mutations in quantitative traits (polygenes, micromutations) It may be pointed out here that mutants classified under the headings of 4-6 are not at all rare. In fact, recent analyses by YAMAGATA, KAWAI, YAMAGUCHI, OKA and others indicate that with appropriate
22 - Hereditas 55
338
A K E GUSTAFSSON AND LVAR GADD
screening methods such mutants can be isolated to a fairly high proportion. Still, there is a considerable dispute concerning the plant breeding value of mutants displaying conspicuous versizs inconspicuous changes (macro- versus micromutations) . The present authors conclude that the procedures to be followed in mutation induction and the screening methods used for the isolation of productive mutants depend on the aims and resources of the plant breeder. The induction of mutations is an additional tool of considerable future value, especially with regard to the isolation of fungus, bacterial and virus resistant mutants, where large M, and M, progenies can be tested by artificial inoculations. But also with regard to properties like increased earliness, lodging resistance and adaptability to fertilizers the mutation method will turn out useful. The techniques of testing the plant breeding value ought to be improved by modern laboratory procedures. An increase in variability by means of induced mutation is easily achieved, both with regard to chromosomal rearrangements, highly productive in the homozygous state, and morphological and physiological changes. Parallel studies in other crop plants, for instance barley, indicate that features like photoperiod insensitivity and thermoperiod adaptability can easily be induced and will no doubt turn out useful also in a crop plant like rice. I n this connection it ought to be emphasized that there are two stages in the application of the mutation method: ( I ) the direct use of mutations in specific traits of agricultural value, for instance earliness and disease resistance, and (2) the indirect use of such mutations in recombination and transgression work. These stages are not independent of one another but should merge into a stepwise procedure of improvement. A plant breeder wishing to improve his rice varieties has to formulate the aim of his breeding work clearly, to analyse the accessible variability and to judge from the existing potentialities about a possible success by ordinary crossing and recombination procedures :IS compared to the use of induced mutation. Many scientists working in the field of mutation research, also in rice, lack in practical knowledge and handling of the crop plant. And, vice versa, practical plant breeders are often remarkably prejudiced against the use of mutations in crop improvement. Both attitudes, the overemphasized fundanieiital one and the prejudiced practical one, are incorrect. A p l m t breeder h a s to a p p l y ccny method useful for his specific aims. Cultivated rice is a n excellent starting material for fundamental research. Being, in addition, one of the chief crop plants of agriculture it offers great opportunities for a
339
successful application of fundamental research, including mutation induction, in practice.
ADDENDUM
Since this article was written, the authors have been informed (T. KAWAI, in lift.) that Japanese scientists have recently released a rice mutant into practice. The new variety Reimei (in English Dawn or Daybreak) was produced at Fujisaka Branch, Aomori Prefectural Agricultural Experiment Station, Aisaka, Towada City, Aomori-ken (Director: Dr. K. TSUNODA). Seed of the variety Fujiminori was treated with y-rays (Co) in 1959. The mutant was isolated in 1960 and tested for yielding capacity and other characters in 1961-1965. The year of release was 1966. The released high-yielding mutant variety differs from the parent by its conspicuously shortened culm. It is highly resistant to lodging (especially under a heavy dressing of fertilizers like the erectoid mutants in barley) and shows less variation in yield due to year and location than the parent (i.e. it has a more stable yield reaction). In addition, it shows the important feature of cool resistance and has a better germination capacity and seedling growth under cultivation at low temperatures.
Acknowledgements. - The completion of this article has been made possible by economical support from the Swedish Research Councils of Agriculture and Natural Sciences, as well as from the National Institutes of Health, Bethesda, Maryland (GM-08877). The authors owe special gratitude to drs. T. T. CHANG, Los Baiios, N. PARTHASARATHY, Bangkok, and B. SIGURBJ~RNSSON, Vienna, for their helpful criticism and well-informed amendments of the text.
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AEE GUSTAFSSON AND WAR GADD
355
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/LEE
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YAMASAKI.Y., SUWA.T. and MURATA. N . 1963. Pathogenicity of radiation-induced mutants in two important phytopathogens of rice . (pp. 244-245) - Genetics Today . Proc . 11th Int . Congr . Genet., The Hague . Sept . 1963. 1 YAMAURA. A . 1933. Karyologische und embryologische Studien iiber einige BambusArten (Vorlaufige Mitteilung) . - Bot . Mag. Tokyo 47: 551-555 . YAO. S Y., HENDERSON. M . T . and JODON. N . E . 1956. CytOlOgiCal evidence of cryptic structural differentiation in the chromosomes as a cause of sterility in hybrids between Indica and Japonica races of cultivated rice. Orgza satiua. - Abstr . Ann . Meet . Am . SOC . Agron., Cincinnati. Ohio. 1956. p 74. YEA. B. and HENDERSON. M . T. 1963 . Effects from irradiation of rice seed with gamma rays and neutrons on several characteristics of the R, generation . Bot . Bull Acad . Sinica 4:90-102 . YEA. P . S . and HENDERSON. M T 1958. Effects from irradiation of dormant seed of rice with gamma rays and neutrons on several characteristics of the R, gene. Agron., Lafayette. Indiana. p . 52. ration . - Agron . Abstr Am . SOC YOSHIDA. Y . 1962. Theoretical studies on the methodological procedures of radiation breeding . I . New methods in aUtOgamOUS plants following seed irradiation Euphytica 11: 95-111 . ZAMORA. M., LENDVAYOVA. 0. and NADURILLE. R . 1963. 1961 supplement to the International bibliography of rice research . Plant Breed . Abstr . 35. No . 4324 .
. .
Contents
Introduction ............................................................ I Origin and genetics of rice .......................................... 1 General references ................................................ 2 . Taxonomy and phylogeny of cultivated rice ........................ 3 . Polymorphism within cultivated rice .............................. 4 . Gene linkages and duplicate factors ................................ 5 . Mode of reproduction .............................................. 6 . Aims of plant breeding ............................................ I1. Spontaneous mutations .............................................. I11. Mutagens and their action in the first generation ...................... 1. Mutagenic agents ................................................ 2 . Radiosensitivity a t the genotypic level .............................. 3 . Comparison between different types of ionizing radiations ............ 4 . Developmental stage and effects of ionizing radiations .............. 5 . Chemical mutagens ................................................ 6 . Translocations in the M, generation ................................ IV. Induced mutations .................................................. 1 Chlorophyll lethals as test mutations .............................. 2 Sterility and dwarfness ............................................ 3 . Viable mutants of morphological interest ............................ 4 Productive mutations of plant breeding value ........................ 5 . Blast resistance and induced mutations .............................. 6 . Biometry of induced mutations in quantitative traits (polygenes) . . . . 7. Screening methods for the extraction of productive mutations ........ V Summary and conclusions ............................................ Addendum .............................................................. Literature references ....................................................
. . .
273 273 273 274 277 279 280 281 288 292 292 293 296 299 301 303 303 303 309 311 313 321 323 332 336 339 339

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H E R E 55-20

MUTATIONS AND CROP IMPROVEMENT. VII. THE GENUS ORYZA L. (GRAMINEAE)

(Received April 9th, 1966)

I. ORIGIN AND GENETICS OF RICE

GUSTAFSSON AND IVAR GADD

MUTATIONS AND CROP IMPROVEMENT. VII

Fig. 2. Pachytene chromosomes of Oryza sativa (by courtesy of dr. S. K. SEN).

GUSTAFSSON A N D IVAR GADD

MUTATIONS AND CROP IMPROVEMENT. VII

XI. 0. breuiligulata Chev. et Rochr.

I\. 0. australiensis Domin

V. 0. minuta J. S. Presl et C. B. Presl 0. punctata Kotschy ex Steud.

BBCC BBCC CCDD CCDD

Wild; Asia Wild; Africa, Asia Wild; America Wild; America

GUSTAFSSON AND IVAR GADD

MUTATIONS AND CROP IMPROVEMENT. VII

AKE GUSTAFSSON AND IVAR GADD

MUTATIONS AND CROP IMPROVEMENT. VII

GUSTAFSSON AND IVAR GADD

MUTATIONS AND CROP IMPROVEMENT. VII

GUSTAFSSON AND IVAR GADD

MUTATIONS AND CROP IMPROVEMENT. VII

GUSTAFSSON AND IVAR GADD

MUTATIONS AND CROP IMPROVEMENT. V I I

GUSTAFSSON -AND IVAR GADD

11. SPONTANEOUS MUTATIONS

MUTATIONS AND CROP IMPROVEMENT. VII

WKE GUSTAFSSON A N D IVAR GADD

MUTATIONS AND CROP IMPROVEMENT. VII

GUSTAFSSON AND IVAR GADD

111. MUTAGENS AND THEIR ACTION IN THE FIRST GENERATION

MUTATIONS AND CROP IMPROVEMENT. VII

2. Radiosensitivity at the genotypic level

GUSTAFSSON AND IVAR GADD

Fig. 3. Differences in radiosensitivity of varieties in rice (redrawn after FUJII, 1962).

GUSTAFSSON AND IVAR GADD

10-50 30-40 20-30

3. Comparison between different types of ionizing radiations

> > >

3.6X108nth/~ni2 1.9 ,, 5.2 ,, 3.6 ,,

> 1.9 >

MUTATIONS AND CROP IMPROVEMENT. VII

X - r a y s (LOkR) Av. ht. L.3cm

Thermal neutrons (5,2x1d3 nth/cm2) Av. ht. 3,6cm

X -rays (30 kR) Av. ht. 8.9cm

Thermal neutrons (3.6~10'~ nth/cm2) Av. ht. 9,2cm

GUSTAFSSON AND IVAR GADD

4 X 1018nth/cm2 of neutrons in the variety Norin 29. These experiments

> 40 kR > 50 kR > 4 hours

& 28 k R 30 kR 2 hours 1-2 hours

MUTATIONS AND CROP IMPROVEMENT. VII

600 m 800 ,, 1,000 1,200 2,000 ,,

ca. 15,120 ,, 5,400 ,, 5,400 ,, 10,800 ,, 13,680

4.0 % 1.0 % 1.2 % 0.4 % 0.5 %

GUSTAFSSON AND IVAR GADD

10 15 Seed water content

MUTATIONS AND CROP IMPROVEMENT. VII

LD,, for gerinina t'ion

LD,, for seedling height

0 . 3 4 . 5% 2 hours 0.1-0.3 % 2 hours 0.04-0.07 % 5 hours k0.3 % 2 hours 0.1-0.3 % 2 hours

MUTATIONS AND CROP IMPROVEMENT. VII

IV. INDUCED MUTATIONS

GUSTAFSSON AND IVAR GADD

0.981 3.312 0.065** 0.147

6.72 6.77 G.76 6.72 6.60'#* 6.66""