Crosses between Hordeum vulgare L. and H. bulbosum L. II. Elimination of chromosomes in hybrid tissues

Summary In interspecific crosses between diploid and autotetraploid cytotypes of Hordeum vulgare L. (cultivated barley) and H. bulbosum L. (bulbous barley grass), several kinds of hybrid endosperm, vegetative tissue and generative tissue were studied. In all these tissues, mitotic disturbances occurred which resulted in chromosome elimination. The chromosomes of H. bulbosum were eliminated more frequently than those of H. vulgare.Differences between the three types of tissue might have been due to differences between their mitotic rhythms. Differences between various kinds of the same type of tissue were discussed in relation to the ratio of the total number of genomes in embryo, endosperm and maternal tissue, and the ratio of the genomes of the parental species within the hybrid nuclei. For both ratios assumptions as to the genetic value of each genome were taken into account.The cause of the disturbances was probably genic disharmony between the parental genomes.     

Influence of Telomeric Heterochromatin Loss on Growth and Development of the Early Embryo and Endosperm in Triticale

The influence of telomeric heterochromatin on early embryo and endosperm development and the production of aberrant endosperm nuclei were investigated using eight triticale (×Tritocmecale Wittmack) lines homozygous for the presence or absence ol blocks of telomeric heterochromatin from rye chromosome arms 1RL, 2RS, (SRS, and 7RL. The loss of heterochromatin from two chromosomes (6R, 7R) produced a significantly lower percentage of aberrant endosperm nuclei than did the loss from one chromosome (6R). Deletion of a third block had a significant additional effect in only one of four lines. The differences between lines for rales of embryo and endosperm development were significant even within a karyotypc, presumably because of genetic variation. The onset of cellularization of the endosperm tended to obscure the detection of aberrancy, and since it evidently occurred at different times in different lines, may have been largely responsible for the apparent reduction in aberrant nuclei in lines with less heterochromatin.     

An embryological study of a Pisum sativum x Vicia faba cross

Summary Flowers of Pisum sativum plants grown in the field and greenhouse were self-pollinated and pollinated with Vicia faba. Germination of pollen and pollen tube growth were slower in the Pisum x Vicia cross. About 2% fertile ovules in the field and 8% in the greenhouse were obtained from the Pisum x Vicia cross, compared to 26% and 48% for Pisum x Pisum. Development of the Pisum x Vicia embryo was normal in form, though much slower than the Pisum x Pisum embryo, and about the sixth day it began to collapse. The greatest difference between the two types of crosses was observed in the endosperm. While the Pisum x Pisum endosperm nuclei divided normally and endosperm developed throughout the ovule cavity and around the embryo, the Pisum x Vicia endosperm nuclei formed only scattered masses of densely stained nuclei which were not always in close proximity to the hybrid embryo. It was concluded that faillure of some ovules to develop following Pisum sativum x Vicia faba cross pollination was the result of slow germination of pollen and slow growth of the Vicia pollen tubes, with subsequent lack of fertilization. Collapse of fertilized ovules was associated with abnormal and limited development of the hybrid endosperm, possibly leading to lack of nourishment and eventual collapse of the hybrid embryo.     

Pollination systems, hybridization barriers and meiotic chromosome behaviour in Nemesia hybrids

The ability of 13 Nemesia species (six annual and seven perennial) to sexually hybridize was investigated. Six of the perennial Nemesia species investigated were inter-fertile with one another. Two of the annual species, N. macroceras and N. strumosa, were inter-fertile. Thirty three crosses were successful and resulted in viable seeds. The analysis of meiotic chromosome behaviour in interspecific hybrids indicated that Nemesia chromosomes in different parental species were homeologous. No evidence of chromosome inversions or chromosome translocations was observed during meiosis in interspecific hybrids between the six perennial Nemesia species. In the hybrids produced between N. macroceras and N. strumosa, a quadrivalent was observed during meiotic metaphase I, indicating that these two species differ by a reciprocal translocation. A successful hybridization was made between N. anisocarpa (annual) and N. foetans (perennial), producing two triploid hybrids. In the unsuccessful crosses, pollen tubes were observed entering ovaries and ovules, suggesting that post-fertilization barriers were preventing sexual hybridization. Many of these crosses produced nonviable, shrunken, empty seeds, suggesting that endosperm breakdown and embryo abortion prevent interspecific hybridization in unsuccessful crosses. The manipulation of ploidy levels in N. fruticans and N. strumosa and tissue culture of N. strumosa × N. fruticans ovules failed to overcome post-fertilization barriers between these species.     

Early Embryo-Sac Development and Intraspecific Variation in the Mean Endosperm Cell Cycle Time in Vicia narbonensis (L.)

The early endosperm development in seven Vicia narbonensis populations was studied under a 23/ 16 ° C day/night temperature regime. Double fertilization occurred in less than 24 h after pollination. Endosperm divisions were synchronous until the 8th division of the endosperm nuclei. Aberrant endosperm nuclei were first observed alter the 8th division and their number increased after the 10th division. Endosperm nuclei continued to divide until the 12th day after pollination. During the following days a decrease in the number of endosperm nuclei was observed together with a substantial increase in the number of aberrants. Endosperm nuclei finally disintegrated. Neither cellularization nor starch deposition was observed in the endosperm. A positive correlation (r 0.69) was recorded between early pod length and the number of endosperm nuclei found in the ovules within each pod. In the populations studied the number of endosperm nuclei increased at a steady but different rate. This was followed by embryo development as well. It was concluded that intraspecific variation in the mean endosperm cell cycle time exists in Vicia narbonensis and that populations A 174 and A 202 should be more compatible to Vicia faba ev. ‘Polycarpe’ than population A 201.     

Interspecific hybridization between Vicia faba (L.) and Vicia narbonensis (L.): Early pod growth and embryo-sac development

Summary Fertilized embryo-sac development and pod growth was studied in one Vicia faba cultivar, one Vicia narbonensis population and their reciprocal crosses. The initial development of endosperm and embryo was at least four days faster in V. narbonensis than in V. faba. Pods and ovules developed also faster in V. narbonensis than in V. faba. The growth rate of the hybrid pods followed the growth rate of the mother species, but was slower than that of the pods from selfed flowers. In the cross V. narbonensis × V. faba the ovules stopped growing 9 days after pollination, while in the reciprocal cross they stopped growing 15 days after pollination. Hybrid embryo-sacs from V. faba × V. narbonensis were aborted before they reached the stage of 256 endosperm nuclei or 200 embryo cells. Selfed V. faba embryo-sacs reached this stage in less than 9 days after pollination. In the reciprocal cross the embryo-sacs were aborted before they reached the stage of 128 endosperm nuclei or 80 embryo cells. Selfed V. narbonensis embryo-sacs reached this stage at the 4th day after pollination. Given that at these stages the embryo has less than 200 cells it was concluded that an in-ovule embryo culture technique should be developed to obtain hybrid plants.     

Study on lily introgression breeding using allotriploids as maternal parents in interploid hybridizations

Based on a recent hypothesis, “Five same genomes of endosperm are essential for its development in Lilium”, it is expected that allotriploid lily (OTO) can be hybridized with diploid Oriental lily (OO) for introgression breeding in Lilium L.. To test the hypothesis, OTO lilies, ‘Belladonna’, ‘Candy Club’ and ‘Travatore’, were used as the maternal parents and crossed with two diploid OO cultivars, ‘Siberia’ and ‘Sorbonne’, and the species L. regale Wilson (TT). Results showed that capsules of all OTO × OO hybridizations developed well and 0.8~3.3 viable seedlings per ovary were obtained through normal pollination and embryo rescue; however, all OTO × TT crosses failed. Genomic in situ hybridization showed that the progenies of the OTO × OO hybridizations were aneuploid and a variable number of T-genome chromosomes were introduced into the progenies through the allotriploid lilies. The present results not only demonstrate that allotriploid OTO lilies, although male sterile, can be used as maternal parents to produce aneuploid progenies, but also strongly support the new hypothesis in lily breeding.     

Inter-specific hybrids of Lilium lankongense Franchet produced by embryo-culture

Summary Lilium lankongense crossed with some related species of Lilium produced seeds without endosperm and with very small embryos which would not germinate in soil. Hybrid plants have been obtained from a number of these crosses by culturing excised embryos on sterile nutrient medium. It is suggested that the technique may prove useful in the production of hybrids from other lily species-combinations which are not normally viable.     

Chromosome association and pollen fertility in some interspecific hybrids of Lilium

Summary Meiotic observations in PMCs were made in eight kinds of interspecific hybrids in Lilium. Three hybrids, 6134-S × L. cernuum. L. regale × L. leichtlinii maximowiczii and L. auratum platyphyllum × L. henryi showed respectively the mean chromosome association frequencies of 1.4_II, 0.8_II and 1.5_II per cell at MI which were the lowest values hitherto reported in Lilium hybrids. In L. longiflorum × L. henryi, 33.3° of the cells had 1 or 2 multivalents of three to five chromosomes in addition to uni- and bivalents. 12° bivalents and 12 univalents were invariably observed in the triploid hybrid L. longiflorum × L. cernuum. The hybrids between the species belonging to the different sections of the genus generally showed high pollen sterility, with some exceptions. A more or less remote genomic homology was found between the different sections in Lilium.     

Intraspecific Variation in Mean Endosperm Cell Cycle Time in Vicia faba (L.) and Interspecific Hybridization with Vicia narbonensis (L.)

The rate of early embryo-sac development was studied in seven faba bean (Vicia faba L.) cultivars grown in a controlled environment. The embryo-sac stage was determined from the number of endosperm nuclei per embryo-sac and the number of embryo cells during the first 12 days after pollination. Differences in early embryo-sac development were observed between the genotypes studied. In addition, five different V. faba×V. narbonensis crosses and reciprocals were made using genotypes with small differences in mean endosperm cell cycle time and genotypes with large differences. The percentage of success (pod set) in V. faba×V. narbonensis crosses ranged between 9 % and 59 % while in the reciprocals it ranged from 12 % up to 30 %. However, cytological studies showed that the high percentage of success (pod set) observed in the cross A-107 × A-202 was not associated with a higher percentage of interspecific hybrid embryos. The results indicate that genotypes of the two species with smaller differences in mean endosperm cell cycle time result neither in a higher percentage pod set nor in bigger hybrid embryos.     

Analysis of endosperm development of allotriploid × diploid/tetraploid crosses in <Emphasis Type="Italic">Lilium</Emphasis>

Lilium are Fritillaria-type plants. Triploid lilies, regardless of their male sterility, can be used as female parents to cross with appropriate males, in contrast to other triploid Polygonum-type plants, which are usually seedless. Up to now, however, little attention has been paid to the reason. Here we made allotriploid × diploid/tetraploid crosses in Lilium. Endosperm and progenies of LAA × AA/AAAA crosses were analyzed for ploidy level and genome composition. The endosperm of some LAA × AA/AAAA ovules developed well but there was little or no development of LAA × LL/OO endosperm. The endosperm genome composition of LAA × AA, LAA × AAAA, LAA × LL, LAA × OO are theoretically derived as 5A + 2L, 6A + 2L, 4A + 3L, 4A + 2L + O, respectively. Genomic in situ hybridization showed that the progenies of LAA × AA/AAAA were aneuploid. Based on the experimental results and analyses, our hypothesis is that five same genomes of endosperms are essential for its development in triploid × diploid/tetraploid crosses of Lilium. The hypothesis can explain the success or failure of 3x × 2x/4x crosses in Lilium and is of importance for lily breeders who need to know the likelihood of success when producing new cultivars of this economically important horticultural crop.     

Problems in breeding and cytology of sugar cane

In connection with the increase of chromosome number found in species hybrids of Saccharum the author studied embryo-sac development within several Saccharum froms from early stages of meiosis in the embryo-sac mother cells to stages where fertilization already had taken place.Normally by meiosis within E M C's a row of four haploid megaspores is formed from which the innermost, the chalazal megaspore, develops into an embryo-sac and the other three degenerate.Haploid embryo-sacs, thus formed, may occur in Saccharum. Frequently however, in Saccharum endo-duplication takes place, that is splitting of the chromosomes within the nucleus of the chalazal megaspore, through which the chromosome number is increased.It is described how within Saccharum by this phenomenon embryo-sacs may be formed with diploid and tetraploid chromosome numbers and with numbers between haploid and diploid.The increase of chromosome number, observed in the young megaspore after meiosis, could be confirmed by counting the chromosomes of dividing nuclei within young embryo-sacs.Chromosome numbers found in dividing zygotes and endosperm nuclei within fertilised embryo-sacs also pointed to the occurrence of egg-cells and polar nuclei that must be haploid, diploid and tetraploid.The significance of this research in relation to plant breeding will be discussed in Part V.     

Genomic Affinity Relationships in the Genus Avena

Many interspecific hybrids between diploid, tetraploid and hexaploid Avena species were obtained without artificial embryo cultures. Most of the hybrids showed variable chromosome pairing with univalents, bivalents and multiple associations, even in PMC in the same hybrid, at first metaphase; making ii very difficult to understand the degree of genome affinity between parent species, and especially between different species. The basic or theoretical chromosome pairing consisting of bivalents and univalents is presented, in which the chromosome ratios (%) statistically agree with observations. In addition, the per cent frequency of chromosomes forming multivalents is calculated. On the bases of these numerical data, the genomic affinity relationships are discussed for present and previously reported interspecific hybrids or Avena.     

普通玉米子粒性状四类效应的遗传分析

采用二倍体胚和三倍体胚乳种子遗传模型及其分析方法,以５个玉米自交系及其间配制的Ｆ１,Ｆ２,ＢＣ１,ＢＣ２为材料,研究了５个玉米种子性状的胚直接效应、胚乳直接效应、母体效应和细胞质效应。结果表明,除粒宽外,各性状的遗传同时由质效应和胚、胚乳、母体基因效应所控制,丰粒重主要受胚乳和母体效应的影响,粒长的遗传以母体效应为主,粒宽和粒厚以胚乳效应为主。各部位子粒百粒重的胚乳直接加性效应与母体加性效应的协方差达显著或极显著水平,其余性状的胚、胚乳直接效应与母体效应间的协方差均不显著,通过母体植株的遗传表现可对这些性状进行有效的选择。在提高丰粒重的玉米育种中,以任一部位子粒百粒重作为选择标准可起到相同的选择效果,直接选择粒宽的分离类型有利于获得大粒的遗传材料。Ｓ２２是改良百粒重的优良亲本。     

普通玉米主要营养品质性状的遗传效应分析

用13个普通玉米自交系按NCⅡ设计（7×6）研究了普通玉米主要营养品质性状的遗传效应。结果表明，普通玉米主要营养品质性状的遗传受种子二倍体胚、三倍体胚乳、细胞质和二倍体母体植株4套遗传体系的控制；主要品质性状4套遗传体系的效应值之间无太大的差异，蛋白质、淀粉和赖氨酸含量的胚遗传效应全部体现为胚加性遗传效应，而4种品质性状的胚乳遗传效应则全部由胚乳显性遗传效应构成；母体遗传效应主要表现为母体加性效应；遗传效应预测值表明亲本9636、99S2041-1-1-1和478在普通玉米品质改良中可能有一定的应用潜力。结果还发现，不同品质性状间存在不同程度的遗传相关，因此对这些性状进行间接选择或杂种优势利用均可取得一定的效果。     

The use of endosperm balance number for predicting gamete selection in complex polyploid interspecific Trifolium repens × T. nigrescens crosses

Endosperm balance number (EBN) was used to predict the outcome of crosses among a series of complex polyploid interspecific hybrids between Trifolium repens and T. nigrescens. Normal endosperm development, and hence seed‐set, requires a 2 : 1 maternal : paternal EBN ratio in endosperm. Based on their crossability, T. repens (2n = 4x = 32) and T. nigrescens (2n = 2x = 16) were assigned the same EBN of 4. This number was used to predict the relative success of different backcrosses and intercrosses. The method was further developed to successfully predict which gametes would effect fertilization when complex polyploid hybrids with uneven genomic constitutions (5x, 7x) produced mixtures of euploid and aneuploid gametes. This was achieved by allocating EBNs to all potential gametes of complex polyploids. Allocation of gamete EBNs enabled not only prediction of the success or failure of a cross, but also which gametes would be filtered to effect fertilization in complex crosses. This enhanced ability to understand the potential outcomes of complex crosses will increase the efficiency of plant breeding programmes that integrate secondary gene‐pools into economic species by interspecific hybridization.     

海岛棉半配生殖品系有性过程的胚胎学观察

半配生殖品系ＶＳｇ自交或以ＶＳｇ作母本与海岛棉标准系３－７９杂交，花粉能在柱头上萌发，花粉管能在花柱组织中生长并能进入胚珠到达胚囊。精核与极核融合、初生胚乳核分裂，以及胚乳游离核形成均表现正常，但精卵融合不同于３－７９。突出表现在一部分胚囊中，进入卵细胞的精核迟迟不与卵核融合，未融合的二核（♀＋♂）“合子”经过一段比３－７９稍长些的“休眠期”后，各自独立分裂，形成各种类型的嵌合胚。     

Production and cytogenetics of tetraploid–octoploid Avena hybrids

The objectives of this research were to gather hybridization information about crosses between tetraploid and synthetically derived octoploid lines of Avena and their reciprocals, and to study the cytology of the hybrids obtained. When the octoploid plants were used as females, only 2.6% of the pollinated florets set seed, while seed set increased to 39.0% in reciprocal crosses. However, germination was 95.7% for the first group of hybrids and only 18.2% for the reciprocals. These diverse results can be explained by the paternal: maternal endosperm genomic ratios and the hypothesis of polar nuclei activation. The tetraploid-octoploid hybrids did not always contain 42 chromosomes. Octoploid lines derived indirectly from tetraploid (AABB) × hexaploid (AACCDD) crosses were found to retain their A and B genome chromosomes. Univalents in the above hybrids averaged 15.44 in 1990 and 15.15 in 1991, indicating the presence of C and D genome chromosomes inherited from the octoploid lines. Directly synthesized octoploids from diploid (A_cA_c) × hexaploid (AACCDD) crosses were found to form 20.44 univalents, 5.30 bivalents, and 2.69 trivalents when crossed with AABB tetraploid species. The high number of univalents indicates that chromosomes from at least three genomes, i.e. B, C, and D, remain unpaired, while trivalents are the result of homology between the A and A_c genomes. Hybrids between tetraploid and octoploid lines can be obtained at low frequencies but, regardless of the direction of the cross, these hybrids are highly sterile.     

Wheat embryogenesis and haploid production in wheat × maize hybrids

Summary Embryogenesis was analyzed in wheat × maize hybrids using paraffin sectioning. Embryogenesis in wheat × maize hybrids is different from that in self-pollinated wheat plants. Development of the embryo is not accompanied by the formation of an endosperm. The endosperm nuclei remain free in the cytoplasm, fail to advance into the cellular stage, and degenerate at a later time. The antipodal cells quickly degenerate in the fertilized ovaries of wheat × maize hybrids similar to self-pollinated ovaries. The antipodal cells remain normal in unpollinated ovaries. The pre-embryo will abort if it is allowed to develop on the plant, because of a nutritional shortage in the absence of an endosperm. Therefore, embryo rescue is necessary for haploid production from a wheat × maize hybrids. Haploid polyembryos were obtained from spikelet culture of wheat × maize hybrids. The formation of polyembryos is due to the cleavage of the pre-embryo and the effect of 2,4-D. The frequency of haploid embryo production and plant regeneration is affected significantly by maize genotypes, but not by wheat genotypes. The concentration of 2,4-D affects only the size of the embryo.     

基于单粒观察值的胚乳性状QTL图的构建方法

谷物胚乳性状是决定谷物品质的一类重要性状。由于胚乳是由雌性的两个极核和雄性的一个精核受精发育而形成的三倍体组织,因此胚乳性状的遗传基础要比通常的二倍体农艺性状复杂得多。本文根据三倍体胚乳性状的数量遗传模型,提出利用分布于染色体上的所有分子标记对胚乳性状数量基因座位（QTL）进行区间作图的极大似然估计方