Cytology of inbreds and f 1 hybrids of pearl millet

Summary The meiotic behaviour of chromosomes was studied in four inbred lines and their F ₁ hybrids of P. typhoides. The inbred lines showed a decrease in mean chiasma frequency when compared with the population plants, but differed from one another in their mean chiasma frequencies. Together with the decrease in mean chiasma frequency the inbreds showed variation in mean chiasma frequencies. The inbred lines showed a number of meiotic abnormalities such as extra chromosomes, extra fragments, desynapsis, persistent nucleoli and differential condensation of chromosomes. The F ₁ hybrids of these inbreds exhibited heterosis for chiasma frequency. All the F ₁'s had mean chiasma frequencies higher than the means of the respective participating parents. The F ₁'s, however, differed in the degree of heterosis exhibited. In the F ₁ hybrids, the variation in mean chiasma frequency, both between plants and between PMC's within plants, was significantly lower than that of the inbred lines. The effect of environment was studied in the inbred lines and their F ₁ hybrids. The mean chiasma frequencies of the inbred lines were significantly lower, and the variation in mean chiasma frequencies was greater, in the stress season. The mean chiasma frequencies of F ₁'s did not show any significant differences between the two seasons. The F ₁'s exhibited less variation in mean chiasma frequency than the inbred lines, showing that F ₁'s were developmentally more stable. The F ₁'s did not show any meiotic abnormalities in either season.     

Genetic and environmental components of chiasma control

A disruptive selection programme for high and low chiasma frequency over four generations has produced two lines of Schistocerca gregaria with mean values of 21.6 and 18.0, respectively. The mean of the base population was 20.4. Realised heritability estimates for both the high and low lines were 27.4% and 48.8% respectively; only the latter value is significant. The non-significant value in the high line has resulted either from unidirectional dominance or because the genes responsible for high chiasma values themselves undergo a higher recombination rate in high frequency individuals and thus reduce the response to selection. — It is suggested that the genes governing chiasma frequency can be regarded as mainly neutral due to the large additive genetic component, at least in low chiasma frequency types. Accordingly, chiasma frequency variation and its adaptive significance may not be such an important component of fitness as has been proposed in the past.     

Genetic interactions between wheat and rye genomes in triticale

Summary Six primary triticale lines were produced from two advanced breeding lines of Triticum durum and three inbred genotypes of Secale cereale. The wheat and rye parents and the triticale derivatives were crossed in all possible combinations within each species group. Chiasma and univalent frequency of parents and hybrids were determined. The primary triticale lines had more univalents and less chiasmata per pollen mother cell than the corresponding wheat and rye parents together. The parental wheat F₁ exhibited negative heterosis for chiasma frequency whereas all rye hybrids had much higher chiasma frequencies than their inbred parents. Triticale F₁s generally showed lower chiasma frequencies and more univalents than their parents, but the degree of pairing failure was dependent upon which of the parental species within the triticale, wheat or rye, was in the heterozygous state. F₁s with heterozygous wheat genome only showed the least reduction in chiasma number (presumably caused by gene actions within the wheat genome), while F₁s with heterozygous rye genome showed high reduction in chiasma frequency and an increase in pairing failure (induced by negative interactions between the heterozygous rye and the wheat genome in triticale). A high correlation was found between the frequency of undisturbed pollen mother cells and the frequency of aneuploids in the subsequent generation. A higher number of aneuploids occurred in those populations which were heterozygous for the rye genome.     

Cytogenetics of Lolium perenne

Summary A number of inbred lines of Lolium perenne have been developed as far as the fifth inbred generation and are being used for basic studies on the cytogenetics of the species. An analysis on variation in the pattern of chiasma frequency and distribution, including the parent plants and all generations down to the S5, reveals that the effect of inbreeding is to reduce chiasma frequency and to increase both the cell and bivalent variances. Evidence is presented for a genetic basis of polygenic control of chiasma formation and distribution, and a model is suggested for control over the three related components of variation which involves a two-stage level of regulation.     

Relationships between frequency,localization and errors in chiasma formation in desynaptic rye

Four inbred lines of rye (Secale cereale) and the F₁ and F₂ from the cross between two of them have been studied. The results indicate that the genotypes used show variation in chiasma frequency, chiasma error frequency and chiasma localization. Significant correlations between these characters have been found: as chiasma frequency decreases both chiasma error frequency and distal localization increase. These correlations lead us to the assumption that these anomalies are in fact secondary effects of the failure of some preconditions for exchange. It is suggested that one such exchange precondition may be effective pairing.     

Genetic and environmental components of chiasma control

Variation in chiasma frequency within and between individuals has been investigated in Schistocerca gregaria and Stethophyma grossum. By taking sequential samples of the same testis in S. gregaria it has been demonstrated that there is considerable variation in chiasma frequency between times, within individuals, and that this variation is not evenly distributed between individuals of the same population. The response and recovery patterns after exposure to successive temperature treatments also indicate a differential sensitivity of individuals within the experimental population. — An analysis of interfollicular variation in chiasma frequency has revealed significant differences between follicles within individuals both in S. gregaria and a Spanish population of S. grossum. A comparable analysis on individuals of S. grossum carrying supernumerary segments shows that the presence of such segments increases the amount of variation between follicles within individuals and also between individuals within the population. — The distribution of cell chiasma frequencies in S. gregaria is normal whereas in Stethophyma cell frequencies approximate to a Poisson distribution. The structurally different supernumerary segments, present in Austrian and Spanish populations of S. grossum, both increase mean chiasma frequency and they modify the between cell variance in different ways — the former interchromosomally and the latter intrachromosomally. — The differences in chiasma frequency between follicles and between times within individuals and the differential reaction to heat shock reflect differences in the pattern of genotype-environment interaction. Similarly it can be argued that the presence of polymorphisms involving supernumerary material may play an equivalent role by increasing the between cell or between bivalent variance within individuals, a unique form of genotype-environment interaction at the endophenotypic level. — Such variation in genotype sensitivity to environmental modification has important adaptive value especially in organisms which are subject to recurrent, often random, changes in their environment.     

Genotype-dependent effect of B-chromosomes on chiasma frequency in Eyprepocnemis plorans (Acrididae: Orthoptera)

In order to analyze the effect of B-chromosomes on chiasma frequency, the offspring of different females of the grasshopper Eyprepocnemis plorans have been studied. From the comparison between individuals of the different families having a given number of B's and between individuals with different numbers of B-chromosomes within the same family, it can be concluded that the effect of the genetic background on chiasma frequency is greater than that produced by the presence of B's, and that there is a between-families variation in the effect of B-chromosomes which, in addition, is negatively correlated with the chiasma frequency of related individuals without B's. This genotype-dependent effect of B-chromosomes is discussed.     

The supernumerary segment system of Stethophyma

Three species of the genus Stethophyma have been cytologically examined and all three show variation both for supernumerary heterochromatic segments and for the distribution of standard heterochromatin among the autosomes. The European species, S. grossum, for example, shows considerable interpopulation variation for standard heterochromatin while two of the populations, from Spain and Austria, show supernumerary segment polymorphism. The segments are located interstitially on the S₁₁ chromosome but occupy different positions in the different populations. — In all species, the presence of the extra heterochromatic segments increases the mean chiasma frequency. Moreover, the influence of the segments upon mean chiasma frequency is different in different populations and in different species. In the Spanish population, the increase is both intra- and interchromosomal whereas in Austria the influence of the segment is completely interchromosomal. — In the American species, S. gracile and S. lineatum, where supernumerary heterochromatic segments are carried on both S₁₀ and S₁₁ chromosomes, the effect on chiasma frequency shows a dosage relationship, an increase in the number of segments per individual being correlated with an increase in mean chiasma frequency. It is suggested that the interstitial segments found in all species have originated by direct duplication of chromosome material. By contrast the terminal segments in S. lineatum and S. gracile may be derived by translocation from a B-chromosome since such a chromosome has been found in one individual of the former species. — The variation in segment structure and the distribution of standard heterochromatin, among the European species of S. grossum suggests that these systems have evolved independently in different populations.On educational leave from the Forest Research Laboratory, Fredericton, N. B. Canada.     

Fertility and meiotic chromosome behaviour in autotetraploid pearl millet

Summary Tetraploidy was induced in outbred pearl millet and selection for high and low seed set was started in the C₁ generation. Segregation in the C₃ generation was observed for fertility and also for meiotic features: per cent seed set in selfed earhead, chiasma frequency, chromosome association and chromosome distribution in pollen mother cells were all affected. However, variation in seed set was observed even between samples not differing in meiotic features. It is apparent that factors regulating seed set in autotetraploid pearl millet were genic as well as chromosomal.A high frequency of univalents and trivalents was the main cause of sterility; quadrivalent misdisjunction was not a significant factor. As univalency decreased with increased chiasma formation, the gain was in the form of quadrivalents. However, individuals not differing in chiasma frequency did differ in chromosome association frequencies, indicating that the dependence of chromosome pairing behaviour on chiasmata was subject to genotypic influence.     

An effect of potassium on chiasma frequency and recombination

High and low concentrations of potassium and calcium were combined factorially and applied to the plant,Lolium temulentum. An effect on mean plant chiasma frequency was detected when such plants underwent meiosis at 30°C. Potassium rather than calcium was shown to be the mineral responsible for the observed effects. High concentration of potassium resulted in an increase in mean chiasma frequency. At 20°C little effect of either mineral could be demonstrated, although potassium again was responsible for most of the variation produced.Potassium has also an effect on stability. At 30°C high concentrations of potassium reduce plant variation, whereas at 20°C the converse occurs and an increase in plant variation results. It is suggested that such a difference between the two temperatures supports the conclusion that different genetic systems control the plant means and plant variation.The effect of potassium on chiasma frequency was confirmed inDrosophila melanogaster by studying its effects on recombination in the X chromosome. A striking influence on body size, possibly correlated with recombination, was also observed.The conflict of such results with previous work in which calcium has been shown to be the most important mineral is discussed. The difficulties of arguing a direct role at meiosis for external treatments is emphasised.     

Polymorphism involving heterochromatic segments in Metrioptera brachyptera

A complex pattern of polymorphism involving terminal heterochromatic segments on L₃ and L₄ chromosomes has been uncovered in eight populations of Metrioptera brachyptera. There are individuals in every population which carry reduced segments on one or both L₄'s. In six populations, enlarged heterochromatic segments have been encountered on the L₃ chromosomes in some individuals. The L₄ system is almost certainly stable although the frequency of L₄ karyotypes does not conform to a Hardy-Weinberg distribution in all populations. Stability of the L₃ polymorphism could not be ascertained. A reduction of L₄ heterochromatin leads to a significant rise both in mean cell chiasma frequency and between cell variance. The effect on chiasma frequency is transchromosomal. The normal pattern of strict chiasma localisation tends to be disrupted in germ lines which include modified L₄ chromosomes. There is a reduction in the number of proximal and distal chiasmata and an increase in the frequency of interstitial ones. It is proposed that the standard L₄ heterochromatin may function in conserving heterozygosity and promoting uniformity between parent and offspring. Partial removal may lead to an effective increase in recombination and produce a greater diversity of genotypes for selection to act upon.     

A correction for allele frequency estimates derived from isofemale lines

Isofemale lines are commonly used inDrosophila and other genera for the purpose of assaying genetic variation. Isofemale lines can be kept in the laboratory for many generations before genetic work is carried out, and permit the confirmation of newly discovered alleles. A problem not realized by many workers is that the commonly used estimate of allele frequency from these lines is biased. This estimation bias occurs at all times after the first laboratory generation, regardless of whether single individuals or pooled samples are used in each well of an electrophoretic gel. This bias can potentially affect the estimation of population genetic parameters, and in the case of rare allele analysis it can cause gross overestimates of gene flow. This paper provides a correction for allele frequency estimates derived from isofemale lines for any time after the lines are established in the laboratory. When pooled samples are used, this estimator performs better than the standard estimator at all times after the first generation. The estimator is also insensitive to multiple inseminations. After the lines have drifted oneN _e generations, multiple inseminations actually make the new estimator perform better than it does in singly inseminated females. Simulations show that estimates made using either estimator after the lines have drifted to fixation have a much greater error associated with their use than do those estimates made earlier in time using the correction. In general it is better to use corrected estimates of gene frequency soon after lines are established than to use uncorrected estimates made after the first laboratory generation. This work was supported by an NSERC fellowship to A.D.L.     

Genetic studies of corn (Zea mays L.) anther culture response

Summary Anthers of two maize (Zea mays L.) inbred lines, DBTS (P₁) and B73 (P₂), their F₁, F₂ and first backcross generations — F₁ x DBTS (B₁), and F₁ x B73 (B₂) — were float cultured in YP medium to study the inheritance of corn anther culturability using generation mean analysis. Significant effects of generation were observed for the three traits measured: anther response (%), frequency of embryos (%) and anther productivity. Variation among the generations was similar for anther response and frequency of embryos: no significant differences were found among the P₁, F₁, F₂ and B₁ means, but the means of P₂ and B₂ were significantly lower than those of the other generations. For anther productivity, the F₂ generation tended to have a slightly higher tendency for multiple embryo formation. A simple additive-dominance model was adequate in explaining the inheritance of anther response and frequency of embryos, but digenic epistasis (additive x dominance) was involved in the inheritance of anther productivity. Additive genetic variance was higher than non-additive genetic variance for all the traits; however, only environmental variance was significant. Narrow-sense heritability estimates were 65% and 75% for anther response and frequency of embryos, respectively. Significant inter-plant variation was observed within generations, even for the inbred line DBTS, but isozymic analysis involving five enzyme loci did not reveal any genotypic variability within the inbred lines DBTS and B73.     

The variability of mating signals and preferences in the brown planthopper,Nilaparvata lugens (Homoptera: Delphacidae)

Male and female brown planthoppers, Nilaparvata lugens(Stål) (Homoptera: Delphacidae), exchange substrate-transmitted signals prior to mating. The pulse repetition frequency of the male song is known to be involved in mate recognition and also to vary among geographical populations. Here the variability of male signals, female signals, and female preferences has been examined within a population. Female preference variation has been partitioned into variation in mean preference and variation in the window of preference of individuals. The genetic component of variation has been examined using isofemale lines. Male signal variation was limited (CV=8%) and was mainly within individuals. Female signal variation was greater (CV=15%). Female mean preference varied little (CV=10%) and was closely matched to the male signal mean, but the preference window was wide (> 4 male signal standard deviations on average) and variable (CV=56%). There was evidence for genetic variation only for preference window. These results are discussed in relation to theories of signal system evolution.     

Genetic Variability in the Progeny of Androgenic Dihaploid Plants and Selection of High Agronomic Performing Lines in Brassica Juncea

Androgenic lines of Brassica juncea cv. PR-45 raised by anther culture, were screen for genetic variation. 393 androgenic plants were transferred to pots to study the R₀ generation. These plants showed substantial variation for different characters. Seed progenies of 27 lines of R₀ plants were sown in the field to study R₁ generation. Androgenic plants within lines were significantly homogeneous for the various agronomic characters studied. Two lines were shorter (18 – 20 %) than the control plants, with a remarkable feature of early maturation. Three lines showed 27 – 31 % higher yield than the parent cultivar.     

Seasonal variation of chiasma frequency in Phyllodactylus marmoratus (Gray) (Gekkonidae — Reptilia)

A study of the male meiotic system in two populations of the gekko Phyllodactylus marmoratus (Gray) has shown that both total and interstitial chiasma frequencies vary cyclically throughout the year. This variation is consistent in each population and was observed over a number of years. The total chiasma frequency (an index of the number of terminal chiasmata) has a different form of cyclic variation than does the interstitial chiasma frequency, and it is argued that they are under independent genetic controls. Reproductive studies suggest that only the sperm with the lowest total chiasma frequencies and greatest range of interstitial frequencies are used for fertilization. An experimental approach has shown that prolonged exposure to low temperature produces a significant increase in total chiasma frequency. It is believed that this environmental cue is responsible for the cyclic nature of total chiasma frequency.     

Genetic variation and population structure in Korean populations of eurya japonica (Theaceae)

We investigated levels of genetic diversity, population genetic structure, and gene flow in Eurya japonica, a widespread and broad-leaved evergreen dioecious tree native to Japan, China, Taiwan, and the southern and southwestern coast of the Korean Peninsula. Starch-gel electrophoresis was conducted on leaves collected from 1,000 plants in 20 Korean populations. All 12 loci examined were polymorphic in at least one population, and the mean number of alleles per locus was 3.79. In addition, mean observed population heterozygosity (Ho_p = 0.425), expected heterozygosity (He_p = 0.462), and total genetic diversity (H_T = 0.496) were substantially higher than average values for species with similar life history traits. Although significant differences in allele frequency were detected between populations at all loci (P < 0.001), <7% of the genetic variation was found among populations (F_ST = 0.069). There was a significant negative correlation between genetic identity and distance between populations (r = -0.341; P < 0.05), but this explained only a small amount of the diversity among populations. Indirect estimates of the number of migrants per generation (Nm) (3.37, calculated from F_ST; 3.74, calculated from the mean frequency of eight private alleles) indicate that gene flow is extensive among Korean populations of E. japonica. Factors contributing to the high levels of genetic diversity found within populations of E. japonica include large and contiguous populations, obligating outcrossing (dioecious plant), high fecundity, and long generation time. Occasional seed dispersal by humans and pollen movement by domesticated honey bees may further enhance gene flow within the species.     

Aging and chiasma frequency in Eyprepocnemis plorans (Orthoptera: Acrididae)

The mean frequency of chiasmata was analysed in 29 males of the grasshopper Eyprepocnemis plorans at three different ages considered representative of the whole adult life in this species. In the factors which could potentially affect the parameter studied—namely age, family, individual and the presence or absence of B chromosomes—significant differences were only observed at the individual level. Aging produces no general effect but interacts strongly with the individuals though the age-induced variations in chiasma frequency are not directional. The fact that the range of chiasma frequency variation found within individuals at different ages is comparable to that existing among individuals indicates that the adaptive role of such variations in natural populations must be taken with caution.     

The influence of parental genotype on the chromosome behaviour of Hordeum vulgare X H. bulbosum diploid hybrids

Summary The PMCs of 74 diploid hybrids involving ten H. vulgare varieties and three H. bulbosum lines were analysed at metaphase I and chromosome number and chiasma frequency recorded. There were differences between parental combinations and between plants within those combinations for both chromosome and chiasma number. It is suggested that these characters are controlled by both parents and that differences between plants within families reflect the heterozygosity of the H. bulbosum parents. Chromosomally stable, high pairing lines have been identified for use in a backcrossing programme to introduce H. bulbosum characters to the H. vulgare germplasm.     

Genetics of speciation in the Aedes (Stegomyia) scutellaris group (Diptera: Culicidae) V. Chromosomal relationships among five species

A comparison was made of karyotypes of 5 species in the Aedes scutellaris group and their hybrids. All species had 3 distinct pairs of metacentric chromosomes (2n=6). These were of similar lengths in Ae: malayensis and Ae. alcasidi, and in Ae. polynesiensis and Ae. pseudoscutellaris. However, chromosome 1 in Ae. polynesiensis and Ae. kesseli, and chromosome 2 in Ae. pseudoscutellaris and Ae. kesseli were of unequal lengths. Meiotic analyses revealed that chromosome asynapsis was frequently seen in species hybrids. There was a significant variation in chiasma frequencies between species and their hybrids. However, the mean chiasma frequency was species specific. In addition, the mean chiasma frequency of species hybrids and the extent of chromosomal asynapsis provided a measure of genetic homology between species. Based on the assumption that a dicentric bridge and an acentric fragment were due to a single crossover within a paracentric inversion loop the following conclusions are made. Ae. malayensis and Ae. alcasidi are polymorphic for one paracentric inversion in chromosome 1. Ae. polynesiensis and Ae. pseudoscutellaris, and Ae. pseudoscutellaris and Ae. kesseli are fixed for one paracentric inversion in chromosome 2. Similarly, Ae. polynesiensis and Ae. kesseli are fixed for one paracentric inversion in chromosome 1. These chromosomal differences between species are discussed with respect to hybrid fertility data reported earlier.