FERTILIZATION DYNAMICS AND PARENTAL EFFECTS UPON FRUIT DEVELOPMENT IN RAPHANUS RAPHANISTRUM: CONSEQUENCES FOR SEED SIZE VARIATION

Seed weight varies significantly within and among fruits of wild radish (Raphanus raphanistrum). To determine sources of this variation, we studied fertilization and seed development following controlled pollinations. Within fruits, central ovules were fertilized prior to distal ovules and attained greater seed size. Ninety-seven percent of the variation in mean seed wt per fruit was explained by an analysis of variance incorporating parental effects, pollination date, and the number of seeds per fruit. We document strong maternal effects on the number of ovules per ovary, the number of fertilized ovules per ovary, the number of seeds per fruit, and mean individual seed wt per fruit. Across females, pollen donor had a slight but significant effect on seed wt; no paternal effects on fertilization rate, zygote number, or seed number per fruit were detected. Within females, with one exception, pollen donor had no significant effect on these components of seed development. Stronger maternal main effects may be due to donor x recipient interactions, cytoplasmic factors, the genetic inequity within triploid endosperm, and/or strict maternal control over resource allocation. The large maternal effects relative to paternal effects may limit the rate at which natural selection acts on paternal traits expressed prior to seed maturation.     

From flowering to dispersal: reproductive ecology of an endemic plant,Astragalus australis var. olympicus (Fabaceae)

Astragalus australis var. olympicus is an endemic plant of the Olympic Mountains, Washington. It is considered a Species of Concern by the U.S. Fish and Wildlife Service. This study focused on the reproductive biology of the plant from flower production through seed germination to identify possible weak points that might contribute to its rarity and impede its conservation. Most plants produced a large mean number of flowers and ovules (314.8 and 4106, respectively), but relatively few of these formed fruits and seeds (25.8 and 3.8%, respectively). In decreasing importance, ovules in fruits were lost to predation, seed abortion, and lack of fertilization. The percentages of these fates differed among sites and years. Excluding pollinators by bagging flowers reduced fruit set by ∼50%, but seed set per fruit and seed mass were unaffected. Germination was affected by scarification, temperature, and moisture availability. About 11% of seeds damaged by predispersal seed predators (weevil larvae) remained viable and were released from dormancy. I hypothesize that predispersal seed predation (over 80% at one site) has a negative effect on population growth. Conservation of this species could benefit from improved fruit set and decreased seed predation.     

POLLINATOR EFFECTIVENESS AND SEED SET IN POPULATIONS OF AGALINIS STRICTIFOLIA (SCROPHULARIACEAE)

Examining variation in pollinator effectiveness and seed production resulting from single pollinator visits can provide a deeper understanding of how pollinators may influence reproduction in plant populations. When comparing populations, differences in the number of seeds produced from single pollinator visits to flowers may not always be attributable to differences in pollen deposition, but rather to differences in plant fecundity or resource availability. Pollinator effectiveness and seed production were studied for two populations over a 4-year period and were measured using single bee visit manipulations of flowers. No significant difference in pollinator effectiveness (pollen deposited on stigmas) was observed between the two populations. However, a significant difference between the two populations was observed in the number of seeds produced per flower. The Wellhouse population produced approximately three times as many seeds/flower from a single pollinator visit as did the Firefly Meadow population. Within each population, pollinators (Bombus pennsylvanicus and Apis mellifera) did not differ in the number of pollen grains deposited on stigmas or seeds produced per flower from single visit experiments. Differences in plant density, pollen viability, and ovules per flower also could not account for a significant amount of the variation. A resource augmentation experiment (water and fertilizer application) resulted in a decrease in seeds per flower per bee visit for the water treatment at the Wellhouse population only. For both populations, pollen deposition, pollen viability, and ovules per flower were unaffected by the resource augmentation. Alternative possibilities for the observed differences in seeds per flower per bee visit are discussed.     

EFFECTS OF POLLEN AND RESOURCES ON SEED NUMBER AND OTHER FITNESS COMPONENTS IN AMELANCHIER ARBOREA (ROSACEAE: MALOIDEAE)

Most Amelanchier arborea flowers have 10 ovules, but the number of filled (embryo-containing) seeds per fruit is usually less than 10 and is highly variable within each individual plant. Because fruit developmental time correlates with seed number, this variation in seed number results in asynchronous fruit ripening. Field experiments tested whether seed number was pollen- or resource-limited. Manipulation of resources in the shoot at the time of fruit initiation by defoliation, girdling, fruit thinning, or foliar feeding had no significant effect on seed number per fruit, although fruit set and seed weight were affected. Supplemental cross-pollination also had no demonstrable effect on seed number. Most ovules that do not become filled seeds are visible as small “undeveloped seeds”; these are not necessarily aborted seeds as this was also the fate of ovules in unpollinated carpels. Alternative hypotheses for the determination of seed number are proposed and discussed.     

Patterns of fruit and seed production inBauhinia ungulata (Leguminosae)

Patterns of seed and fruit production ofBauhinia ungulata, a small tree legume indigenous in tropical America, were studied in Costa Rica. Only about 8% of flowers produced fruits. The average pod had 19 ovules and about two thirds of these began seed development, with mature pods containing an average of 9.7 mature undamaged seeds. About half of the mature pods were damaged by herbivores and within these, 27% of ovules or seeds had been eaten. Among trees there was no significant variation in pod production, but the number of ovules per pod and seed production per pod varied significantly. Within infructescences most pods were retained at middle positions. Within pods, the probability of an ovule developing into a seed increased toward the distal end. The pattern of seed and fruit production in this species agrees well in general with that reported for other neotropical legumes. The abortion of seeds and fruits can be regarded as a way of controlling maternal investment, and as a response to herbivory.     

A TEST OF SEVERAL HYPOTHESES FOR THE DETERMINATION OF SEED NUMBER IN AMELANCHIER ARBOREA,USING SIMULATED PROBABILITY DISTRIBUTIONS TO EVALUATE DATA

The number of seeds per fruit is variable within Amelanchier arborea trees. Because A. arborea flowers are five-carpellate and each carpel contains two ovules, we were able to use the pattern of seed maturation within fruits to test whether the failure of some ovules to develop into seeds is determined by mechanisms operating at the level of carpels, such as stigma-clogging, or by mechanisms operating at the level of ovules, such as ovule infertility. The presence of one-, two-, and zero-seeded carpels demonstrated that the number of ovules developing into seeds was not due entirely to carpel-level phenomena. In order to test the hypothesis of carpel independent seed development, without the assumption that all ovules have the same probability of developing into seeds, it was necessary to use simulation, since no conventional statistical models were appropriate. Analysis of this simulation allowed us to reject carpel independent processes as the only determinant of seed number. A mixed model of seed development, in which some carpels fail entirely and ovules in the remaining carpels develop equiprobably, was shown to be consistent with the data.     

Application of a probabilistic model for analysing the abortion of seeds and pods in winter oilseed rape (Brassica napus)

In winter oilseed rape (WOSR), only a subset of ovules can develop into seeds in the majority of pods. Any difficulty during the process of seed production may result in seed or pod abortion. This study aimed to reproduce the process of seed development in WOSR based on a limited number of parameters. As a result of the complexity of the developmental patterns of WOSR, it is challenging to identify the roles of various factors that influence seed production using an experimental approach. Here, we present a stochastic probabilistic model of seed development. The generalised least squares method was implemented to estimate the model parameters using the experimental data. Experiments were done in Grignon (France) in 2008 and 2009. The variations in the parameters were analysed according to the following four factors: year, pod rank, inflorescence position and ramification‐clipping treatment. The year had no effect on the number of ovules per ovary (μ) and the probability of seed viability (p). The proportion of effective pollen grains (k) significantly decreased with pod rank at the end of the main stem. Inflorescence position influenced the number of ovules per ovary (μ: 30.8–33.8 from top to bottom) and the parameter k. The mean number of seeds per pod on the main stem and the bottom ramification along the stem was larger than the other ramifications within one plant. Ramification‐clipping treatment increased the number of ovules per ovary (μ: 31 for control plants and 32 for clipped plants), the parameter k and the number of seeds per pod (p). This effect could be due to the competition for assimilates between the pods and seeds. Furthermore, the distribution parameters of the pollen number per stigma (m) remained stable, and the probability of pod survival (Bo) varied with different factors, including the year, pod rank and inflorescence position. Our results indicate that pollen germination is a factor that determines final seed number. This model can identify the impact of each of the factors that lead to the abortion of seeds and pods in WOSR, which include the position effect, assimilate competition and pollination limitation. However, further studies on the pollination process in WOSR should help to refine this model.     

Predispersal seed predation on five Piper species in tropical rainforest

Absolute number of seeds lost to predispersal seed predators and proportion of total seeds lost per infructescence were compared among five Costa Rican Piper species of different annual fecundities. Mean seed number and mean seed size in the five species were negatively correlated. The impact of predation on these species was inversely related to the number of seeds they produced. The two early successional species had very high fecundities, a combination of many seeds per infructescence, many infructescences per plant, and, in one species, year-round reproduction. Although seed predators destroyed as many or more seeds of these early successional species than they did of the less fecund, late successional species, this loss accounted for a relatively minor proportion (9 and 12%) of the seeds of the early successional species. In contrast, late successional species produced fewer, larger seeds in a smaller number of infructescences and were not continually in fruit. One of these species, which produced intermediate numbers of intermediately sized seeds, lost 30% of the seeds in each infructescence on average. Seed predators destroyed a larger proportion (65 and 76%) of the seeds per infructescence in the two species with fewest seeds per infructescence. High levels of insect damage in these late successional species caused many of their infructescences to abort prematurely. Taken together these factors resulted in annual fecundities several orders of magnitude smaller in shade-tolerant Piper species than the annual fecundities of shade-intolerant, early successional species. Seedlings of the two early successional species were common in large gaps and other sunny clearings and seedlings of the species with 30% seed loss were occasional, whereas no seedlings were seen of the two species with the highest proportional seed loss, suggesting that seed predation on the latter species may limit seedling recruitment.     

Density-dependent variation in reproductive success in a terrestrial orchid

 Within population variation in plant density can affect reproductive success and breeding systems. We examined such effects in Listera cordata R. Br., a north temperate terrestrial orchid, in a subalpine population in Colorado, USA. Hand pollinations showed that L. cordata was self-compatible as all pollinations produced fruits. Cross-pollinations differed significantly from self-pollinations, and had a higher number of seeds per capsule and higher % of fertilized ovules. Together they could compensate for the transmission advantage of self-pollinations. Average number of flowers per inflorescence was 4.2–4.4 and average fruit set was 20.7–23.7%. By establishing 36 plots with varying plant densities (1, 4, 8 and 16) and emasculating all flowers in target plants of a set of 20 plots, we estimated density effects on fruit and seed production. Plant density had no effect on the amount of reproduction except in reduced seed production of highly dispersed plants. Our results suggest that inbreeding depression may be an important selective factor in L. cordata. Further studies are needed to define its role in the evolution of mating systems in this species. Received February 21, 2000 Accepted December 26, 2000     

Size-dependent reproduction in Australian alpine Ranunculus

Abstract The effect of plant size on reproduction in four species of alpine Ranunculus (R. muelleri, R. dissectifolius, R. graniticola and R. niphophilus) was investigated in two sites over two seasons in the field on a total of 190 plants. The effects of plant size (number of leaves) and number of flowers on the number of anthers, ovules and seed per flower and per plant were determined. There was a positive relationship between several measures of reproduction and plant size in all four species, indicating that reproduction is size-dependent. All the results indicate that the main factor controlling the amount of seed produced by alpine Ranunculus is the size of the plant. Specifically, bigger plants produced more seed by producing more flowers, not by producing more ovules per flower, or higher seed set per flower. Correspondingly, bigger plants produced more anthers by producing more flowers, rather than by producing flowers with more anthers. The total number of seeds produced by a plant was directly proportional to plant size in the four species. Therefore, reproductive effort should not vary with plant size in the four species.     

Variation of pollination and resource limitation in a low seed-set tree, Liriodendron chinense (Magnoliaceae)

Pollen limitation and resource limitation have been documented as the major factors responsible for plants commonly producing more ovules than seeds, but few studies have examined pollen deposition directly in natural populations at different sites and times. We investigated the causes of low seed set in four populations of Liriodendron chinense (Magnoliaceae), an insect‐pollinated endangered tree endemic to southern China, over 2–3 years. One pistil potentially produces two ovules. The number of pistils per flower varies among populations, but in three of the four populations the variation in a given population was not significantly different among years. Overall, populations with higher pistil numbers tend to set more seeds per flower, but a positive correlation between pistil numbers and seed production per flower was observed in only one of the four populations. The numbers of pollen grains deposited per stigma varied from 0 to 60. The proportion of pollinated stigmas per flower ranged from 44% to 88% among populations and years. The numbers of pollen grains deposited per stigma and the percentages of pollinated stigmas were significantly different between populations, and two populations showed significant differences between years. A positive correlation between stigmatic pollen load and seed set was sought in ten population‐by‐year combinations but, in a given population, high stigmatic pollen loads did not always result in high seed set. Examination of pollen deposition, pistil and seed production over several sites and years showed that in addition to pollination, other factors such as resource or genetic loads were likely to limit the (lower than 10%) seed set in L. chinense. It appears that small, isolated populations experience severe pollination limitation; one population studied had seed/ovule ratios of 0.84% and 1.88% in 1995 and 1996. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society, 2002, 140 , 31–38.     

Reproductive biology ofCistus ladanifer (Cistaceae)

The phenology, major floral characteristics, fruiting levels, and breeding system ofCistus ladanifer L. (Cistaceae), a common western Mediterranean shrub species, were studied in a southern Spanish population. The white, large (64 mm in diameter) flowers of this shrub appear during spring (March–May) and produce abundant pollen and nectar. In the year of study, flowers lasted up to three days, during which they were visited by a diverse array of insects including beetles, flies, and bees. Hand-pollinations revealed that flowers do not set any seed unless cross pollen is applied to the stigma. Microscopical observations indicate that self pollen tubes grow down the stigma but invariably fail to induce fruit maturation. At the plant level, all estimates of fecundity investigated (number of seeds per capsule, proportion of ovules developing into seed, and proportion of flowers setting fruit) were highly dependent on nearest neighbour distance, with isolated plants setting as little as 0% fruit. In contrast, plants within a clump often transformed into fruit as much as 90% of the flowers. At the population level, seed output was estimated to range between 3,000 and 270,000 seeds per plant during 1991.     

Sexual reproduction and variation in floral morphology in an ephemeral vernal lily,Eyythronium americanum

Summary In a riparian population of Erythronium americanum (Liliaceae) in central New Jersey, experimentally self-pollinated plant produced markedly fewer fruit and fewer seeds per fruit than hand-outcrossed and open pollinated plants, even though differences were not evident between pollen tubes that penetrated stigmas from self or foreign pollen. This weak self-compatibility and a positive relation between the percentage of seeds set by outcrossed plants and the distance between pollen donor and recipient plants indicate that this population could be susceptible to inbreeding depression.Limited resources for seed development apparently constrained maximal seed production, based on low seed set (40.6%) by hand-pollinated plants and positive correlations for these plants between plant size and the number and size of seeds set. In contrast, naturally-pollinated plants set a smaller proportion of their ovules, suggesting that limited pollinator service reduced the quantity of seeds produced in this population. Free-foraging bees usually removed more than half of the available pollen in a single visit, so that individual plants probably have few opportunities to disseminate their pollen.Even though sexually reproductive ramets produce only a single flower per year, less than a third of variation in floral morphology is associated with variation in plant size. Within the flower, the sizes of some closely associated structures, such as the style and ovary, and the anthers and filaments, vary essentially independently of one another. Production of nectar and pollen, the ultimate attractors of pollinating insects, was positively correlated with flower size.     

Factors affecting intrafruit pattern of ovule abortion and seed production in Hormathophylla spinosa (Cruciferae)

We investigated the pattern of intrafruit seed production in Hormathophylla spinosa (Cruciferae) over a 7-yr period. H. spinosa fruit have two chambers, each chamber containing two ovules, and usually situate perpendicular to the infructescence axis with one chamber above the other. The percentage of ovules ripening to seeds is usually lower than 50%. In addition, we found a consistent position-dependence in seed ripening probability, since more than 90% of the ripe seeds are arranged in the lower chamber of the fruit. This pattern of seed production is not produced by the direct or indirect effect of seed predators, by pollen limitation or by nonrandom fertilizations. By contrast, fruit removal experiments showed that sibling rivalry can partially explain the seed production pattern, be the cause of the reduction in the seed number from the uppermost threshold of 50%. Moreover, experimental manipulations of fruit orientation suggest that the position-dependence in seed production is the cause but not the effect of seed ripening. We presume that some architectural effect is producing a significant increase in the ripening probability of the ovules arranged in the lower chamber, causing thus the observed pattern in the intrafruit seed production in H. spinosa and severely constraining the maximum number of seeds per fruit to two.     

Variation in the costs and benefits of mutualism: the interaction between yuccas and yucca moths

Summary Yucca moths are both obligate pollinators and obligate seed predators of yuccas. I measured the costs and net benefits per fruit arising for eight species of yuccas from their interaction with the yucca moth Tegeticula yuccasella. Yucca moths decrease the production of viable seeds as a result of oviposition by adults and feeding by larvae. Oviposition through the ovary wall caused 2.3–28.6% of ovules per locule to fail to develop, leaving fruit with constrictions, and overall, 0.6–6.6% of ovules per fruit were lost to oviposition by yucca moths. Individual yucca moth larvae ate 18.0–43.6% of the ovules in a locule. However, because of the number of larvae per fruit and the proportion of viable seeds, yucca moth larvae consumed only 0.0–13.6% of potentially viable ovules per fruit. Given both oviposition and feeding effects, yucca moths decreased viable seed production by 0.6–19.5%. The ratio of costs to (gross) benefits varied from 0% to 30%, indicating that up to 30% of the benefits available to yuccas are subsequently lost to yucca moths. The costs are both lower and more variable than in a similar pollinator-seed predator mutualism involving figs and fig wasps.There were differences between species of yuccas in the costs of associating with yucca moths. Yuccas with baccate fruit experienced lower costs than species with capsular fruit. There were also differences in costs between populations within species and high variation in costs between fruit within populations. High variability was the result of no yucca moth larvae being present in over 50% of the fruit in some populations, while other fruit produced up to 24 larvae. I present hypotheses explaining both the absence and high numbers of larvae per fruit.     

REPRODUCTIVE PROFICIENCY IN PAEONIA CALIFORNICA (PAEONIACEAE)

Study of three populations of geophytic Paeonia californica in southern California revealed considerable variation, within and between populations, in both potential and in realized seed output. Plants are weakly self-compatible, but natural seed set (x = 4.2 seeds per carpel) is closer to seed set produced in hand cross-pollinations (x = 4.5) than in self pollinations (x = 1.4). Flowers are protogynous, nectariferous, and scented; 17 species of bees in four families, and one species of melyrid beetle were found in them and probably account for pollinations at these sites. Counts of reproductive parts show greatest inter-population variation in mean numbers per plant, of flowers (population means ranged from 16.2 to 21.2), ovules (744 to 1142), pollen grains (23 million to 38 million), and % viable pollen (26.6 to 81.4). Seed set also varied significantly, with only 11.4 to 26.8 % of the ovules maturing into seeds. This low reproductive success is probably correlated with the genetic anomalies studied by others—particularly the translocation heterozygosity that causes lowered production of viable gametes. The relatively few seeds produced are heavy (x = 0.184 g per seed), and germination percentages are high (x = 80%). The elongation of cotyledon bases during hypogeal germination seems to permit optimal germination at soil depths of only 2–3 cm, but environmental factors such as drought stress and herbivore pressure promote high seedling mortality.     

Plasticity of reproductive components at different stages of development in the annual plant Thlaspi arvense L.

Summary This study examines the effect of different densities and the removal of all neighbours at different stages of development on all components of reproduction in the inbreeding annual Thlaspi arvense L. A 64-fold increase in density significantly reduced all repooductive components. The number of flower buds per plant was decreased most strongly; the order of decreasing plasticity among the other components was number of capsules per flower, individual seed weight, ovule number per capsule, flowers per flower bud and seeds per ovule. Removing neighbours at all stages of development increased seed yield of plants in comparison to the control without density reduction, but patterns of plasticity depended strongly on time of treatment. The main effect of the removal of neighbours at the vegetative stage was to increase the number of flowers per plant, but the number of ovules per capsule and seed weight increased also, and abortion of capsules decreased. Removing neighbours at the onset of flowering initially failed to affect flower number per plant, instead it resulted in a strong reduction of capsule abortion and an increase in seed weight. However, several weeks after flowering had initially ceased, fresh lateral inflorescences were produced, resulting in a second flush of reproduction. Removing neighbours at the stage of fruit ripening resulted at first only in an increase in seed witht, but later a second reproductive phase occurred. Fresh lateral branches were produced, but the apical meristem was also reactivated. The overall pattern of plasticity among all reproductive components in response to a removal of neighbours was the same as in response to density. The position of a capsule along the inflorescence influenced its number of ovules, the rate of seed abortion and the mean weight of seeds, with the type of effect depending on the developmental stage at which neighbours were removed. Significant negative correlations were found between the mean weight of seeds and the number of seeds in a capsule under all treatments.     

Breeding system and seasonal variation in seed set in a population of Potentilla palustris

The breeding system and factors affecting seed set in a population of Potentilla palustris were studied in Central Jutland, Denmark. The reddish-brown, bowl-shaped, and nectariferous flowers of Potentilla palustris had an average of 376795 viable pollen grains and 187 single-ovuled carpels. The number of ovules per flower decreased significantly along an inflorescence, from 312 in the basal flowers to 97 in the flowers in distal positions. The flowers were protandrous and opened mainly during the day. Male and female phases lasted on average 1.6 and 3.0 days, respectively. Pollination experiments showed that P. palustris was facultatively autogamous to facultatively xenogamous. The plant was selfcompatible and auto-deposition of pollen occurred, but insect polllination enhanced seed set. Outcrossing increased seed set compared to selfing. A long outcrossing distance increased seed set comparative to short distance crossings. Seed set per flower declined through the flowering season mainly due to a significant reduction in the number of ovules per flower with time and/or node position. It is suggested that intra-plant competition for a limited amount of resource is the main factor limiting seed set in this population, but pollen limitation occur in late season. The quantitative and qualitative impact on seed set by pollinating insects is discussed.     

EFFECTS OF ENVIRONMENTAL STRESS ON COMPONENTS OF REPRODUCTION IN CLARKIA UNGUICULATA

The effects of stress on reproduction in Clarkia unguiculata were investigated by monitoring flower number, seed number and seed wt after subjecting plants to drought and heavy fruit loads, and by monitoring seed number after pollinating overage flowers. Fewer main-stem flowers were produced in drought stressed plants than in non-stressed plants. Drought stress did not affect overall seed numbers and seed wt in 1981, but did significantly affect seed number and wt in different flower age classes. Overall seed numbers were not affected by drought stress in 1982, but seed wt was significantly lower in plants with stressed ovules. Fewer flowers but more seeds per pollination were produced by plants in which all or ½ the flowers were pollinated than by plants in which ¼ or 1/6 of the flowers were pollinated. Seed wts were lowest in plants with a full fruit load and in fruits produced late in the growth cycle. Seed number was variable but high following pollination of flowers with 1–7 day old stigmas, and much lower in flowers with older stigmas.     

Inheritance of quantitative traits in crosses between two <Emphasis Type="Italic">Pisum sativum</Emphasis> subspecies with particular reference to their breeding value

The experimental study was conducted during the period of 2008–2010 at the experimental field of the Institute of Forage Crops in Pleven. The hybridization scheme included direct and back crosses covering four varieties of forage pea (Pisum sativum L.), namely two spring ones, Usatii 90 and Kamerton from Ukraine, and a winter one from Bulgaria, Pleven 10. There was analyzed the inheritance of quantitative traits such as plant height, height to first pod, pod number per plant, seed number per plant, seed number per pod, seed weight per plant and number of fertile nodes per plant of parental components (P₁ and P₂) and both first (F₁) and second (F₂) hybrid generations. The cross Usatii 90 × Pleven 10 showed the highest real heterosis effect for plant height (8.26%), pods per plant (158.79%), seeds per plant (272.16%), seeds per pod (42.09%), seed weight per plant (432.43%) and number of fertile nodes per plant (117.14%). The cross Pleven 10 × Usatii 90 had the highest real heterosis effect height to first pod (11.06%). In F₂ plants, the strongest depression for plant height (5.88%), seeds per plant (57.88%), seeds per pod (55.93%) and seed weight per plant (55.99%) was in the cross Usatii 90 × Pleven 10, for height to first pod (1.47%) in the cross Kamerton × Pleven 10 and for number of fertile nodes per plant (15.91%) in the cross Pleven 10 × Usatii 90. The highest positive degree of transgression for number of fertile nodes per plant (165.64%) and seed weight per plant (162.10%) was in the cross Pleven 10 × Kamerton and for pod number per plant (102.54%) and seeds per plant (99.13%) in Kamerton × Pleven 10. The stability of the characters was determined. Low variability in F₁ and F₂ was found in plant height (3.97–6.85%). Variability of number seeds per plant in F₁ was highest (11.86–33.23%). For all other traits, the variability varied from average to high. A lower narrow-sense heritability coefficient was observed for plant height, height to first pod, pods per plant, seeds per plant and seed weight per plant (from 0.001 to 0.230). In few cases, such as in fertile nodes per plant (0.39 and 0.81) and seeds per pod (0.44), the coefficients of broad-sense heritability were higher.