The distribution of genes for yellow pigment of flour in a doubled haploid (DH) population obtained by anther culture.
A DH population from the cross `ATS7/L 1063 = S29 (Pro 1 + Pro 2)/S29 (Lr19 + Rht1)' was compared with the F₂ hybrids of the same combination for flour color. It is well known that the gene Lr19 is linked closely to the gene for yellow pigment of flour. The theoretical expected segregation for the DH population was 1 dark yellow:2 yellow:1 light yellow:1 creamy:2 white:1 light white. In fact, the group of plants with cream-colored flour was greater than expected. The DH lines derived from microspore callus showed no groups of plants with dark yellow and yellow flour. However, one DH line, derived from callus, had a light yellow flour. The absence of DH plants with dark yellow and yellow flour, derived from callus, can be explained by selection of microspores without the yellow pigment gene, which is linked with Lr19 in the androgenetic process in vitro or the loss of the yellow-pigment gene in the hemorhizous process.
The creation of a hairy leaf durum wheat for protection against Oulema melanopus L.
For protection against the wheat beetle (Oulema melanopus L.), we performed transfers of the Hl genes (hairy leaf) from T. dicoccum, T. persicum, T. timopheevii, Ag. intermedium, and Ag. trichophorum into the background of the best local cultivars of durum wheat.
The larva of the wheat beetle, in some years, causes essential damage to cultivars with smooth leaves. The isogenic lines for the Hl gene were obtained by the backcross method. The cultivars Bezenchukskaya 139 and Charkovskaya 46 were used as recipients. The trichome length in these isolines was 0.16-0.20 mm. F₁ hybrids from the cross of the durum wheat cultivar Ludmila with Ag. trichophorum, which have hairy leaves (trichome length 0.33-3.52 mm on the flag leaf) were obtained. The flag leaves of the F₁ hybrids have trichomes 0.70-1.87 mm in length. The Hl genes, in this case, have partial dominance.
By crossing the cultivar Ludmila with Ag. intermedium, F₁ hybrids were obtained with two types of leaf hairs. The first type was as in Ludmila, and the second type as in Ag. intermedium. In this case, codominance was observed.
The Moscow People's Friendship University
Ul. Efremova 18, lcv. Y, Moscow 119048, Russia.
A.K. Fedorov.

Ontogenesis of the alternative wheat.
Among agricultural crops, a number of alternative [editors note: facultative] types can be encountered, particularly in wheat, barley, rye, peas, and the perennial forage grasses. The alternative wheats, in contrast to winter and spring types can be seeded in autumn or spring.
As a rule, the varieties of alternative cereals are grown in regions where winter is comparatively mild. They are most widespread in those regions where autumn seeding is during conditions of comparatively short days. On the other hand, the growing of winter varieties is more frequent in those regions where cereals are seeded at an earlier date, under conditions of long days, and at relatively high light intensity. Besides their great importance in agricultural production, the biology of the alternative plants is incompletely studied. Among the problems to be studied are how the alternative varieties differ from the spring varieties and what properties bring about winter hardiness when planted in autumn. Our investigations (Fedorov 1971, 1989) revealed that, in the alternative plants, the influence of the prevailing autumn conditions is much stronger than in the summer crops. Particularly important was the influence of short days, which delay growth and development, thus enabling plants to survive in winter when planted in autumn.
To access the differences in developmental characters between alternative wheats and spring and winter varieties at different seeding dates, vernalized and nonvernalized seeds were sown simultaneously (vernalization period was 50 days). As an example, the condensed data from one of the trials are presented in Table 1.
The experimental data in Table 1 show, when seeded in the spring, alternative wheat varieties will reach the heading phase almost at the same time as spring varieties or with an insignificant delay. The winter varieties did not ear by the end of the vegetation period. When sown in the autumn, plants of the alternative wheats displayed a significant lag in development as compared to the spring varieties. This is concluded from the date of the beginning of differentiation of the growing point (shoot apex). In wheat varieties with greater winter hardiness from the more northern regions (Czech alternative and 26191 from Austria), the slowing down of the developmental processes occurred earlier and to a larger extent than in varieties exhibiting poorer winter hardiness and in those from more southern regions (Bulgaria, Inner-Asiatic Republics). The property of the alternative varieties to exhibit a significant delay in growth and development under the influence of conditions prevailing in autumn, is an adaptive feature enabling them to survive the winter. The slowing down of growth and development observed in the alternative plants when sown in the autumn is caused essentially by short days, poor light intensity, and, in part, by reduced temperatures. The comparatively rapid development of the alternative plants, with shooting and development of inflorescence in the autumn brought about by these and other causes (sowing in the early autumn, supplementary
light, seeding with vernalized seed), unfavorably influence winter hardiness. The alternative varieties must be sown at a date that does not allow them to reach the shooting phase in the autumn, while achieving sufficient development at the same time.
Table 1. The development of wheats depending on seeding dates and seed vernalization.
____________________________________________________________________________________________
Cultivar Seed used Period from full emergence to Period from full emergence to
for sowing differentiation of shoot apex, days full earing, days
Seeding dates
_____________________________________ ________________________
5/1 6/1 6/16 8/1 8/11 8/21 5/1 6/1 6/16 8/1
____________________________________________________________________________________________

Lutescens unvernalized 15 13 11 20 21 28 44 42 34 nv
62 vernalized 13 13 10 19 20 22 42 40 32 nv
Czech unvernalized 28 26 23 75 nv nv 59 57 nv nv
alternative vernalized 18 16 12 27 42 66 46 43 36 nv
26191 unvernalized 28 25 23 76 nv nv 58 57 nv nv
vernalized 19 16 12 27 42 66 44 43 36 nv
NOE unvernalized 26 25 22 72 nv nv 60 58 58 nv
vernalized 16 16 11 23 38 55 45 43 34 nv
109 unvernalized 22 22 19 46 59 nv 50 50 48 nv
vernalized 14 13 10 21 24 25 43 42 32 nv
Surhak unvernalized 24 20 - 47 60 nv 48 47 - nv
5688 vernalized 14 12 - 20 21 22 41 40 - nv
Lutescens unvernalized 109 nv nv nv nv nv nv nv nv nv
329 vernalized 17 16 14 30 46 69 46 45 38 nv
____________________________________________________________________________________________

Note: spring wheat - Lutescens 62; alternative wheats - Czech alternative, 26191, NOE, 109, Surhak 5688;
winter wheat - Lutescens 329.
Vernalization of seed has little influence on development of alternative wheats when sown in the autumn, under the conditions of comparatively short days. Vernalization of the seed results in a significant acceleration of the development of an alternative wheat. Frequently, the greater the delay in a variety under given light conditions, the more development was accelerated by seed vernalization. Under field conditions, alternative wheats were similar to the winter varieties. The vernalization process comes to an end at the onset of the severe winter period. The capacity of the alternative wheats to survive under the influence of lower temperatures is a consequence of vernalization, their relationship to light (i.e., they lag behind in development to a lesser extent) under the influence of the short day, and the lower light intensity. This is particularly important in those regions where the spring begins comparatively early, still in short-day conditions, and where dry spells are frequent (e.g., in the Inner-Asiatic Republics).
Several research workers (Razumov et al. 1961) noted that, in some southern mountainous regions, varieties were encountered with almost similar or even higher requirements for long days compared to the spring varieties of the northern regions that grow under conditions of rather long days. This fact is difficult to explain. Experimental data obtained from the study of alternative plants are helpful in explaining the spread of such varieties in the southern geographical latitudes. To answer this question, we present some data from our experiments with the spring wheat variety, Irody 1006 from central Asia (day length during the growing period generally not longer than 12 hours) and the variety K-190 14 (according to VIR catalogue) from Ethiopia (longest day length of the year is 12 hours and 40 minutes). These two varieties, during a 12-hour day, lag in development almost to the same extent as the spring wheat variety Lutescens 62, when grown in regions where the day length is 17-18 hours. Thus, under short-day conditions, the onset of differentiation of the shoot apex fell in the variety Lutescens 62 on the 26th, in Irody 1006 on the 28th, and in K-19014 on the 28th day from full emergence.
Under similar conditions, when vernalized seed was used, the southern varieties developed ahead of Lutescens 62. Thus, the onset of the differentiation of the shoot apex in Irody 1006 fell on the 16th day after emergence, in K-19014 on the 18th, and in the spring wheat Lutescens 62 on the 21st day. It is evident that both spring wheats from the southern regions, under short day conditions, respond to vernalization to a greater extent than Lutescens 62. This seems to indicate that these southern varieties, with regard to developmental properties, are nearer to the alternative varieties than the more northern variety, Lutescens 62. In our opinion, this is due to the fact that the southern varieties, with regard to the localities where they are grown in the initial phase of development, are often exposed to the influence of lower temperatures. For instance, Irody 1006 is sown not only in the spring, but also in autumn and winter. This variety, when spring sown, often is exposed to the influence of low temperatures, particularly at night. In Ethiopia, the variety K-19014 is grown at an altitude of more than 2,000 m where temperatures are much lower than in the valleys, especially in the night. In some southern plant varieties, an adaptation evolves that causes them to lag behind in development at a day length where they are exposed to the influence of low temperatures or where they are in advance of the occurrence of low temperature.
Therefore, varieties grown in comparatively short day in the southern geographical latitudes lagged behind in development when grown in short-day conditions almost to the same extent as the varieties from northern regions grown under long day conditions. Many plants originating from southern regions (wheat, barley, oat, flax, and peas) exhibit a considerable delay in development under short-day conditions. It is a much longer delay than would be expected by the natural day length in the districts where they are grown. This can be explained essentially by the fact that their habit is, to a certain extent, nearer to that of the alternative plants.
All wheats are long-day plants. There are no photo-neutral wheats. They are distinguished by the degree of development of this property. Varieties originating from regions near the equator express this property poorly, as do varieties originating from the high latitudes farther from the equator.
Under autumn short-day conditions, with low light intensity, lower temperature, i.e., the conditions of autumn sowing, alternatives grow slower compared to spring crops, which increases their winter hardiness. This is more pronounced in the winter hardy varieties than in the less resistant ones.
Thus, in one of our experiments with sowing on August 11, the shoot apex of a Bulgarian alternative differentiated 26 days later than that of the spring wheat Lutescens 62, and the shoot apex of the Czech alternative differentiated 66 days later. The same regularity holds for the relationship to growth intensity. Similar examples are found in the perennial forage alternatives (Timothy grass, alfalfa, clover, and others). This phenomenon can be used in plant breeding as a rough assessment of the winter hardiness of a given plant.
The capacity of alternatives to slow down their development and growth in response to short days and other autumn-like conditions plays approximately the same role in their winter hardiness as vernalization in winter varieties. It retards the formation of reproductive organs and of autumn-shooting, thus bringing about plant adaptation. Alternatives that for some reason form shoots in the autumn form embryonal heads and do not acquire the necessary resistance (winter hardiness).
An early autumn sowing (when the day length is still relatively long) also has a negative effect on the winter hardiness of the alternatives. In this case, they may display embryonal heads and still form shoots in the autumn. In our experiments with wheat sown on August 1, all the alternatives, including such winter hardy varieties as the Czech alternative and K 26191, froze, whereas the winter variety Lutescens 329 showed a resistance of 41 %.
With sowing on September 6 and plant growth and development proceeding on a relatively short-day regime, the alternatives survived as well as the winter wheat Lutescens 329 and even better than Kooperatorka. In this case, the alternatives overwintered, as did the winter varieties, in the tillering phase with a nondifferentiated shoot apex.
When short days (one of the factors retarding the development and growth of alternatives) were artificially prolonged by supplying light during the dark periods, winter hardiness in the alternatives decreased. On August 21, a number of alternative varieties were sown in an illuminated plot. After germination, plants were exposed to different light conditions: 1) continuous illumination, natural plus artificial with normal 500 W incandescent bulbs at an intensity of about 4,000-6,000 lux; 2) normal day; and 3) a short, 9-hour day.
Table 2 shows that light conditions in autumn have a considerable influence on the winter hardiness of alternatives. Complete freezing was observed in the alternatives after continuous illumination, a result of their rapid growth and development under such conditions. Other normal- or short-day regime alternatives wintered as satisfactorily as the winter wheats.
Table 2. The effect of autumn illumination on plant winter hardiness.
_____________________________________________________________________________________

Variety Autumn illumination State of plants before winter period % Wintering
_____________________________________________________________________________________

Lutescens 329 normal tillering, undifferentiated shoot apex 97.5
Czech alternative ` ` 97.1
K 26191 ` ` 94.3
NOE ` ` 79.1
Lutescens 62 ` shooting, differentiated shoot apex 0
Lutescens 329 continuous tillering, undifferentiated shoot apex 98.7
Czech alternative ` shooting, differentiated shoot apex 2.3
K 26191 ` ` 1.9
NOE ` ` 1.4
Lutescens 329 9-hour day tillering, undifferentiated shoot apex 90.5
Czech alternative " " 90.0
K 26191 " " 89.8
_____________________________________________________________________________________

Note: spring wheat - Lutescens 62; alternative wheats - Czech alternative, 26191, NOE, 109, Surhak 5688; winter wheat - Lutescens 329.
It is necessary to time the sowing of the alternatives so that they do not produce shoots before the onset of winter but are able to develop sufficiently. The same holds for perennial grass alternatives.
It was shown above that alternatives, after a prolonged exposure to low temperatures (approximately the same as for the vernalization of winter varieties), display markedly less growth and development on a short-day regime as compared with alternatives not exposed to low temperatures. This may be related to the fact that, in the alternatives, a change of reaction toward short days takes place under the influence of prolonged low temperature (Table 1). This property is of great importance for normal growth and development in the spring. The alternatives are grown mostly in places where spring begins early (when the day is still short) and, if the short-day reaction remained the same as in autumn, their growth and development would be retarded and yield reduced.
Experiments to determine the period required to produce a short-day reaction change revealed that the wheat, NOE requires 35 days and the Czech and 26191 alternatives require 45 to 50 days of low-temperature exposure.
Our experiments to determine length of vernalization demonstrated that alternative and winter varieties, originating from the same geographical region, generally have identical vernalization (in length and conditions). In the field, vernalization begins and ends approximately at the same time. For example, the Czech alternative, alternative 26191, the winter types Lutescens 329 and Mironovskaya 808, and F₁ plants from their crosses have similar vernalization lengths, about 45 days. In the Moscow region, the vernalization of the alternative and winter wheats terminates in October or November, depending on the variety. The normal course on vernalization begins when the average day temperatures are about +10 C or below.
Vernalization is a facultative process that takes place under certain conditions (autumn) and does not take place under others (summer). The role of vernalization in the ontogenesis of alternative plants causes changes in their photoperiod reaction (light reaction). As a result, they lose their ability to delay growth and development considerably.
The length of the vegetative period for spring sown alternative plants is conditioned by the light reaction in the nonvernalized plants (the 1st photoperiodic reaction), but for winter-sown plants, it is conditioned by the light reaction in the vernalized plants (the 2nd photoperiodic reaction).
It was shown, by determining the duration of the reaction to day length, that this period is much longer in alternatives than in spring varieties, lasting 21 days in Lutescens 62, as compared with 45 days in NOE, 26191, and the Czech alternative. In the same varieties sown from vernalized seed, it lasted 24 days, almost one-half the time.
The F₁s of winter and spring crosses, and the F₁s of `alternative wheat x winter wheat', had limited development under short-day conditions, compared to the F<sub>1</sub>s from crosses between spring and winter wheats. F<sub>1</sub>s obtained by crossing alternative wheats headed only at the end of summer. In development type, reaction to light, and winter hardiness, the first generation hybrids were between the parent varieties.
Different crossing combinations of the developmental types differed in the proportion of winter forms observed in F<sub>2</sub>. The F<sub>2</sub> of `spring x alternatives' had winter forms. All plants headed after spring sowing. In the F₂ of `winter x spring' crosses, a small percentage of winter types, 5-10 % segregation, occurred. In the F<sub>2</sub> of `winter x alternatives', quite a large percentage, up to 50 %, occurred. In the development of the alternative wheat type, the reaction to light and winter hardiness are intermediate between those of winter and spring wheats.
The alternative plants, in contrast to the spring forms, under the influence of autumn conditions (short day, poor light intensity) lag significantly in growth and development, which promotes their winter hardiness. This property is more explicit in the alternative varieties originating from more northern regions (e.g., Czech alternative). For plants surviving winter, particularly cereals, a certain adaptability evolves in the course of their phylogeny that delays development under light conditions, enabling them to possibly avoid winter killing.
Such conditions for the alternative wheats are comparatively short days and poor light intensity, because they are usually sown in growing regions under such conditions. For winter plants, such conditions are a comparatively long day and light of sufficient intensity. They are generally seeded in their growing districts under such light conditions (autumn begins there at a relatively early date). In plants able to survive winter under such conditions, a more or less normal yield usually is only possible when vernalization takes place. As a result of the vernalization process, plants surviving the winter change their relationship to light conditions as autumn seedings. Under conditions of longer and more intense light, greater than the normal light intensity to which plants are exposed when sown in autumn, heading is possible even without a previous exposure low temperature. For the alternative plants, such conditions are supplied by the natural light in spring and summer (in the northern and central zones). This is why the alternative varieties have normal yield when seeded in the spring. As evident from our trials (Fedorov 1959), such conditions will arise for winter plants, e.g., at constant light of 50-70 thousand lux.
Differences in the type of plant development (winter, alternative, or spring); mode of life (annual or perennial); and duration of the vegetative period of plants are determined largely by their light reaction at the seedling stage.
The type of plant development, as well as length of the vegetative period, cannot be conditioned by vernalization. It is a facultative process that takes place under certain conditions (autumn) and does not occur under others (summer).
The type of plant development is due to different reactions to light as seedlings (in the Gramineae, at the tillering phase). Spring-type plants are able to slightly delay development under short days. The alternative plants are able to considerably delay under short days, and winter plants can delay development under short and long days.
These results indicate that the characteristics of spring and alternative varieties (delaying development under short-day conditions) and of the winter varieties (delaying development under both long- and short-day conditions) are similar phenomena and are determined by the photoperiodic reaction during tillering. This reaction, and not vernalization, determines the difference in the length of a plant's vegetative period, particularly in hybrids and initial varieties.
The photoperiodic reaction in unvernalized plants is expressed to a slightly different degree from that in vernalized plants. As a result of vernalization, plants lose the ability to adapt, expressed as a lag in development under definite light conditions. Therefore, the light reaction is affected. Plants of all types respond to vernalization with an acceleration of development depending on their reaction to light. They respond only under definite conditions of illumination, and the higher the response, the greater the delay. Thus, in wheat, differences in the type of plant development and the length of the vegetation period are due to their different light reaction in the tillering phase and related response to vernalization.
The length of the vegetative period for spring-sown plants (spring and alternative) is conditioned by the light reaction in the unvernalized plants (we call it the 1st photoperiodic reaction), but for winter-sown plants (alternative, winter), it is conditioned by the light reaction in vernalized plants (we call it the 2nd photoperiodic reaction).
The role of vernalization in the ontogenesis of plants is to change their photoperiod reaction (light reaction). As a result, they lose their ability to delay growth and development considerably under influence of the photoperiod preceding wintering (alternative plants of short photoperiod and winter plants of short and long photoperiods). References.
Fedorov AK. 1973. Some data on genetics of wheat ontogenesis. Proc 4th Int Wheat Genet Symp (Sears ER and Sears LMS eds). University of Missouri, Columbia. Pp. 801-803.
Fedorov AK. 1989. Physiological-genetical basis for the type of plant developmental length of the vegetation period in wheat. Cereal Res Commun 17(2):121-127.
Rasumov VI. 1961. Environment and Development of Plants. -M.-L. Selchozgiz. 368 p.
Russian Academy of Agricultural Sciences
Information and Computation Centre, P.O. Emmaus 171330, Tver, Russia.