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.     

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.     

Single-Seeded Fruits and Seedling Establishment in Dalbergia miscolobium Benth. (Papilionaceae)1

Dalbergia miscolobium (Papilionaceae) is a tree of the Brazilian cerrado that produces ovaries with two ovules. Only one ovule usually matures, but when two mature, they are smaller than seeds of single-seeded pods. Germination percentage was lower in small seeds than in medium or large seeds. In general, small seeds gave rise to smaller seedlings (dry mass) than did medium and large seeds. The root/shoot ratio, however, was very similar for the first six months, indicating that this ratio is consistent across seedling sizes. There was no correlation between seed size and relative growth rate. Single-seeded fruit production in D. miscolobium resulted in greater seed reserves and increased fruit dispersal distance; single-seeded fruits were lighter than double-seeded ones. Together, these two features should increase the probability of successful seedling establishment of seeds from single-seeded versus double-seeded fruits.     

Seed germination patterns of Salvia mellifera in fire-prone environments

Summary I examined the effects of pollen loads containing pollen from one, three and five donors on fruit production and fruit quality in Campsis radicans. Number of pollen donors had no significant effect on % fruit production, seed number, seed weight or seed germination. In singledonor pollinations the identity of the donor did have a strong effect on the above parameters. Furthermore, the best single donor sired fruits with more seeds and heavier seeds than any mixture containing this donor. This pattern indicates interference of pollens or preemption of some ovules by the inferior pollen. In Campsis, therefore, the number of pollen donors contributing to a pollen load is less important than the identity of these donors in determining fruit production and fruit quality. Seeds from fruits resulting from mixed pollination were slightly more variable than seeds from fruits resulting from single-donor pollinations.     

EFFECTS OF NONRANDOM SEED ABORTION ON PROGENY PERFORMANCE IN PHASEOLUS COCCINEUS L

In this study, we sought to determine if Phaseolus coccineus normally aborts potentially viable seeds, and whether seed abortion is nonrandom with respect to progeny vigor. The ovaries of Phaseolus coccineus typically have six linearly arranged ovules. The three ovule positions at the stylar end are more likely to mature seeds than the three ovules at the basal end of the ovary. When we destroyed the developing ovules at the stylar end of the fruit after fertilization but before seed abortion, there was a significant increase in the probability that the ovules in the three basal positions would produce a mature seed. The probability of seed maturation in control fruits (no ovules destroyed) ranged from 38.3 to 42.7% over the three field seasons, whereas in the experimental fruits it ranged from 64.3 to 79.7% (similar to that of ovules at the stylar end in control fruits). We did not find any significant change in the probability of seed production in the three ovule positions in the stylar end of the ovary (the positions with high probability of seed maturation) when the basal ovules were experimentally destroyed. These results indicate that potentially viable seeds are regularly aborted in P. coccineus, with seed abortion more frequent in the basal ovule positions than at the stylar positions. In two greenhouse studies and one field study, we compared the vigor of progeny produced in the control fruits with the vigor of progeny produced in fruits where three ovules (either stylar or basal end) were destroyed. We found that the performance of the progeny from the three positions at the stylar end of the control fruits did not differ from the performance of the progeny from experimental fruits in which the three basal ovule positions were destroyed. In contrast, the progeny from the basal positions of the control fruits outperformed the progeny from the experimental fruits when the seeds were produced in the three basal ovules (stylar ovules destroyed). Our findings indicate that when the experimental treatment increased the probability of seed maturation, there was a significant decrease in the average vigor of the progeny in the remaining (basal) ovule positions.     

EFFECT OF OVULE POSITION ON SEED PRODUCTION,SEED WEIGHT,AND PROGENY PERFORMANCE IN PHASEOLUS COCCINEUS L. (LEGUMINOSAE)

We examined the effect of ovule position within the ovary on the probability of seed maturation, on seed weight, and on progeny performance in the outbreeding legume Phaseolus coccineus. Ovaries of P. coccineus possess six linearly arranged ovules (ovule position one = stylar end). We found that in both 1987 and 1988, ovule position had a significant effect on the probability of seed maturation under field conditions. In 1987, ovule positions one. two, and three had a higher probability of maturing seeds than the three most basal ovule positions. In 1988, the probability of producing a mature seed in ovule position one was more similar to the three most basal ovule positions than to ovule positions two and three. The position of the ovule in the ovary had no significant effect on seed weight in 1987, but it had a significant effect in 1988. Overall, seeds from ovule positions one, two, and three tended to produce heavier seeds than the three most basal ovule positions. The effects of ovule position on progeny performance were determined in a greenhouse and a field study. In the greenhouse study, we found no significant overall effect of the position of the ovule that produced the seed on progeny performance. In the field study, we did find a significant ovule position effect on several measures of reproductive performance as well as an overall effect on reproductive performance. In addition, we found a significant interaction between ovule position and number of seeds per fruit. Progeny from the stylar end of the fruit outperformed the progeny from the peduncular end in fruits containing many seeds, whereas there were no significant differences between progeny produced in the stylar and peduncular ends of fruits containing few seeds. Causes of position effects are unknown but hypotheses abound.     

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.     

Intra-inflorescence variation in floral traits and reproductive success of the hermaphrodite Silene acutifolia

BACKGROUND AND AIMS: Intraspecific variation in floral components and reproductive success is often located at the intra-individual level. The arrangement of flowers within inflorescences may explain a great deal of this variation. The variation in number of ovules, fruit set, number of seeds per fruit, seed set, seed weight and seed germination is investigated at different positions within the inflorescence of Silene acutifolia. METHODS: Data were obtained in natural populations, and germination experiments were conducted in a germination chamber. KEY RESULTS: The number of ovules, fruit set, number of seeds, seed set and seed weight, showed a significant decline from early (primary) position to later (tertiary) position. The patterns of intra-inflorescence variation were consistent in different populations and years of study. Seed germination showed an opposite pattern, seeds from primary position showed the lowest germination percentages and seeds from tertiary position the highest, although the effect of position on germination was only marginally significant. There was significant among-population variation in number of ovules per flower. Fruit set also varied significantly among populations, with lower fruit set in the smaller and more isolated population. No significant among-population differences were detected in number of seeds per fruit and seed set. Seeds from the smallest and more isolated population (Arnado) were the lightest. Seed germination showed strong differences between populations, seeds from Arnado started to germinate later, and showed the lowest final germination percentages. CONCLUSIONS: Architectural effects or resource competition are the most commonly proposed hypothesis to explain these patterns. Data suggest that there is less pollen available to pollinate tertiary flowers, and that there is not enough outcross pollen in Arnado. The germination percentages suggest that there is variation in the source of pollen within inflorescences, with high probability of receiving outcross pollen in flowers from primary position, and higher probability of geitonogamous crosses in tertiary flowers.     

Does successful ovule development depend on its position within the pod? Examples from Neotropical Fabaceae

Previous studies have shown a nonrandom pattern of ovule fate probabilities according to ovule position in legume pods. Here, we tested how ovule position within the pods of two Fabaceae affects its fate. We expected higher proportion of well‐formed seeds near the fruit tips and of unfertilized and aborted ovules near fruit bases. We collected pods of Poincianella pyramidalis and Anandenanthera colubrina in a seasonal dry forest in northeastern Brazil and recorded total pod length and ovule number, position, and fate (unfertilized, well formed, aborted, and predated). The proportion of well‐formed ovules at fruit tips was significantly higher than at fruit bases in P. pyramidalis. The opposite pattern was found for unfertilized and aborted ovules, thus corroborating our hypothesis. However, the probability of seed predation in A. colubrina was significantly higher in pod tips, thus providing moderate support for our hypotheses. Interspecific differences in the patterns of ovule fate are likely to be driven by species pollination systems.     

Selective seed abortion affects the performance of the offspring in Bauhinia ungulata

BACKGROUND AND AIMS: Under the microgametophytic competition hypothesis, a non-random pattern of seed abortion is expected, in which only the most vigorous seeds reach maturity. In a previous study, it was found that Bauhinia ungulata (Fabaceae) exhibits a pattern of seed abortion dependent on the position of the ovule within the ovary; ovules located in the stylar half of the fruit, close to the point of entry of pollen tubes to the ovary, have a low probability of seed abortion, whereas ovules in the basal half of the fruit are aborted with a high probability. METHODS: A series of experimental fruits was generated, in which ovules from either the stylar (treatments 1 and 2) or the basal (treatments 3 and 4) half of fruits were destroyed, to evaluate whether these patterns of selective seed abortion have an effect on the vigour of the offspring in B. ungulata. KEY RESULTS: Only 53 % of the seed from control fruits germinated. Seed set in fruits from treatments 1 and 2 showed a significantly lower (33-43 %) percentage of germination; the germination of seeds from fruits in treatments 3 and 4 (49-51 %) did not differ from control seeds. In addition, it was found that the differences in vigour of the offspring are not random with respect to the position of the ovule in the pod. CONCLUSIONS: The overall performance of the seeds correlated with their likelihood of maturation. Seeds located at the basal half of the treatment fruits showed lower values of vigour than seeds located on the stylar half. The differences were more marked for early measures of fitness.     

Effect of ovule position within the pod on the probability of seed production in Bauhinia ungulata (Fabaceae)

BACKGROUND AND AIMS: It has been claimed that ovules linearly ordered within a fruit differ in their probabilities of reaching maturity. This was investigated by studying the effect the position of an ovule within the pod has on seed abortion and seed production in Bauhinia ungulata. METHODS: Fruits collected during the dry seasons of 1999, 2000 and 2001 were opened, and the number, position and status of each ovule within the fruit were recorded. A GLM model was used to assess the effects of population, tree identity and ovule position within the pod on ovule fertilization, seed abortion, seed damage and seed maturation in two populations of B. ungulata. KEY RESULTS: Nearly 30% of the ovules were not fertilized in 1999; this percentage dropped to 5% the following two years. Seed abortion (50%) and seed damage (15%) were the same every year during the study period. Only 15% of the initial ovules developed into mature seeds in 1999; this value increased to 35% in 2000 and 2001. However, seed survivorship was dependent on the position of the ovule within the pod; non-fertilized and early aborted ovules were found more often near the basal end of the ovary. The frequency of seed damage was not affected by position. Mature seeds were found mainly in the stylar half of fruits, where ovules are likely to be fertilized by fast pollen tubes. CONCLUSIONS: The pattern of seed production in B. ungulata is non-random but is dependent upon the position of the ovule within the pod. The results suggest that the seeds produced within a fruit might differ in their vigour.     

Order of fertilization within the ovary in Phaseolus coccineus L. (Leguminosae)

Summary Phaseolus coccineus typically has six linearly arranged ovules per ovary. The three ovules near the stylar end of the fruit (positions one, two, and three) are more likely to produce mature seeds, to produce heavier seeds, and to produce more vigorous progeny than the ovules in positions near the peduncular/basal end of the fruit (ovule positions four, five, and six). We conducted a series of field experiments designed to supplement our understanding of the mechanisms determining these position effects. We found that approximately 98% of the ovules in 752 fruits were fertilized — about 0.6% of the stylar ovules were not fertilized, whereas 3.2% of the basal ovules were unfertilized. Moreover, we found that only about 49% of the ovules in these 752 fruits produced mature seeds. Over 60% of the stylar ovules produced mature seeds, whereas only 37% of the basal ovules produced mature seeds. Consequently, the proportion of fertilized ovules cannot explain the differences in seed maturation among the ovule positions. We found that after 6.5 h most of the fertilized ovules were located in the stylar ovule positions, and that there were no fertilized ovules in ovule positions five and six, indicating that the stylar ovules are fertilized first. When only the fastest growing pollen tubes were permitted to enter the ovary (due to exision of the style), only the ovules at the stylar end were fertilized, indicating that the ovule positions that are fertilized first are indeed fertilized by the fastest growing pollen tubes.On leave from the Escuela de Biologia, Universidad de Costa Rica, Cuidad Universitaria Rodrigo Facio, San Jose, Costa Rica, Central America     

Diversification of seed arrangement induced by ovule rotation and septum formation in Leguminosae

The plants of Leguminosae, which comprises 731 genera, bear monocarpellary flowers. The fruit, known as a pod, generally contains a row of seeds, and the internal structure appears to be simple. In Leguminosae, the seeds are generally arranged longitudinally along the suture of the pod but, in a few genera, namely, Cassia, Indigofera, and Senna, they are arranged transversely. Transverse seed arrangement facilitates the accommodation of a considerable number of seeds and has been considered to be established by rotation of seeds during development. We studied the development both these patterns of seed arrangement, i.e., longitudinal and transverse, in Senna and Indigofera in order to elucidate the structural changes occurring in ovaries (or pods) and determine the forces responsible for rotation of ovules (or seeds) by dissecting ovaries and pods at different developmental stages. Our results showed that at an early developmental stage the ovules were obliquely oriented or transversely arranged; later, the direction of their growth was restricted by the ovary walls or neighboring ovules, and therefore the obliquely oriented ovules rotated inward or outward. Thus, it was clarified that the processes involved in the formation of the internal structure of pods are regulated in a complex manner. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Structural aspects of ovule and seed development and nonrandom abortion inMelilotus officinalis (Fabaceae)

Summary Only one ovule matures into a seed inMelilotus officinalis. Although eight ovules form within an ovary, only the basal ovule develops into a mature seed, whereas the other ovules degenerate. The investigation of ovule and seed structure at different developmental stages and a comparison of quantitative characters of differently fated ovules within an ovary were undertaken by light, phase contrast, and fluorescence microscopy. In this species, campylotropous ovules develop simultaneously on marginal placentae in an apocarpous unilocular gynoecium. Megasporo- and megagametogenesis proceed normally and are completed in bud. The maturation of the Polygonum type embryo sac takes place after the flower opens. Shortly before fertilization, synergids show signs of degeneration in all ovules. At this stage, neither the structure nor the sizes of ovules within one ovary differ significantly. In spite of this, only the basal ovule develops into a seed. Rarely, one of the upper-situated ovules or the basal and another ovule mature into seeds. Seed enlargement is insignificant until the stage when globular embryo and nuclear endosperm are formed. At the seed-filling stage, other ovules have collapsed and the seed gradually comes to occupy the total volume of the pod. The fruit-to-seed length ratio decreases considerably during seed ripening. At fertilization, ovary length is four times greater than ovule length. In the mature state, the fruit and seed lengths are approximately equal. Seed size and weight diminish with an increase in seed number within a pod, although pod size remains constant. It is assumed that nonrandom abortion of young seeds inM. officinalis is under maternal control and is not related to structural abnormalities in ovule development or with limitation in pollen. We suppose that evolution of this species may have proceeded in the direction of a decrease in seed number and an increase in its sizes, which may play an important role in seed dispersal and seedling establishment.     

两种互逆资源梯度影响下刺叶锦鸡儿的种子选择性败育格局

种子的选择性败育在被子植物中普遍存在,开展结籽格局及其影响因素的研究有助于深入了解种子的形成机制及多样化的生殖对策。以刺叶锦鸡儿Caragana acanthophylla为材料,通过传粉过程、胚珠发育动态和资源分配状况的研究,以探讨种子的选择性败育格局及相关影响因素。结果显示：(1)刺叶锦鸡儿具高度自交不亲和性,为泛化的传粉系统,蜂类是主要的传粉者。自然状态的结实率为(86.00±4.96)%,不存在传粉限制,但受精具明显的时间效应。(2)刺叶锦鸡儿单花期4—5d,每朵花有(14.00±0.14)粒胚珠。胚珠的发育从荚果顶部开始,其中,开花后第3天荚果顶部胚珠开始膨大;第11天绝大多数胚珠出现了膨大,此时,在荚果基部的胚珠开始败育,随后荚果顶端受精后的胚珠也出现败育,最终仅荚果中部形成2—3粒种子。其结籽格局为成熟荚果中部胚珠,而败育荚果顶部和基部胚珠的选择性败育类型。(3)受微地形影响,居群内开花植株具斑块分布,对较少开花植株通过添加水肥进行资源调控后,结籽率显著提高,说明在种子形成过程中存在资源限制。综上所述,受精顺序和资源限制两个互逆的资源梯度决定了刺叶锦鸡儿的结籽格局。其中,...     

Effects of parentage,prior fruit set and pollen load on fruit and seed production in Campanula americana L

Summary We conducted a controlled crossing experiment to examine the effects of maternal and paternal parentage, the size of the pollen load, and prior fruit production on the proportion of flowers that set fruit, seed number per fruit and seed weight in a natural population of Campanula americana. Effects due to the maternal parent were large for all measures of fruit and seed production, while the paternal parent had a significant effect only upon mean seed weight. As the number of prior fruits on the maternal plant increased the probability that a flower would produce a mature fruit, the number of seeds per fruit, and total seed weight per fruit all decreased. We found no effect of the size of the pollen loads used in this study on fruit or seed production. These results are consistent with those of other studies that suggest in natural plant populations maternal effects, especially environmental maternal effects, can have an overwhelming effect on fruit and seed production and on seed characteristics.     

Fruit mass variability in Vaccinium myrtillus as a response to altitude,simulated herbivory and nutrient availability

Variability in fruit mass may respond to environmental conditions, in addition to the genetic effect that has been reported in several reproductive features. We first analyzed fruit traits in 15 localities along an altitudinal gradient (300–2000 m a.s.l.) in the berries of Vaccinium myrtillus. We then tested whether changes in natural conditions, such as nutrient availability and herbivory by ungulates, affected fruit features in this species. Therefore, nitrogen (N) addition and clipping were applied in plots situated in six of the 15 localities. In these plots, we later collected fruits in two consecutive years in order to obtain fruit mass, seed number and initial number of ovules per flower ovary. Data along the altitudinal gradient showed that fruit mass was determined by both ovule number and seed number, although these factors only explained approximately half of the variation. Fruit mass and number of ovules and seeds followed a similar pattern with altitude, increasing until reaching a maximum at around 1700 m. Regarding the effect of the experimental treatments, fertilized non-clipped plants produced the heaviest fruits but clipping alone had no significant effect on fruit mass. Moreover, high soil pH had a positive influence on fruit mass. In addition, whereas the original number of ovules per ovary was not affected by the experimental treatments, the number of seeds per fruit increased and the abortion rate decreased in fertilized non-clipped plots, which means that the effect of simulated herbivory on these fruit traits depended on N addition. In conclusion, fruit mass was clearly affected by nutrient availability and this effect is attributed to an increase in the production of seeds rather than to an increase in module size and number of ovules per ovary. This outcome suggests that fruit mass is a plastic trait influenced by environmental factors like nutrient availability.     

BIRD-DISPERSAL OF PHYTOLACCA AMERICANA L. AND THE INFLUENCE OF FRUIT REMOVAL ON SUBSEQUENT FRUIT DEVELOPMENT

Characteristics of fruits, seeds, and ripening of Phytolacca americana were studied in New Jersey, USA to assess their importance in the dispersal strategy of this species. Removal by birds was directly related to the percent of ripe fruits available on a raceme. Fruit removal from a raceme resulted in lower fruit pulp and seed weights for fruits developing on the same raceme, but these differences probably do not influence probability for dispersal by birds. We demonstrate that characteristics of the fruit pulp may decrease seed availability for the primary seed predator, Peromyscus leucopus, and increase the time fruits are available for dispersal. We speculate on the significance of the sequential fruiting in Phytolacca and the relationship to availability of dispersal agents and high-lipid fruits. Sequential ripening and protection from predation result in seed dispersal from late August to early December.     

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.     

A stochastic model of seed dispersal pattern to assess seed predation by ants in annual dry grasslands

A combined field experiment and modelling approach has been used to provide evidence that ants may be responsible for an observed lower patchiness and higher plant diversity in the neighbourhood of ant nests, within Mediterranean dry grasslands belonging to the phytosociological class Tuberarietea guttatae. The hypothesis was that seeds occurring in clumps may have a higher probability to be harvested than seeds having a scattered distribution. In order to test this hypothesis, four analysis steps were performed. First, pattern of seed production and dispersal of four species was recorded; two of them were more abundant next to ant nests (Tuberaria guttata, Euphorbia exigua), whereas the other two were more abundant away from ant nests (Bromus scoparius and Plantago bellardi). Second, a stochastic model was developed to simulate the observed dispersal patterns of each studied species. Third, 10 seed spatial arrangements in accordance to the distribution patterns created by the model were offered to ants and the location of predated seeds was recorded. Finally, the observed pattern of seed predation was matched to models performed by different distributions of probability. Results showed that the probability of being predated decreased as distance among seeds increased. This preference of ants for high concentration of food items holds down the dominant species sufficiently to allow the subordinates to survive, thus increasing diversity near nests. The observed higher frequency of small-seeded, small-sized, or creeping therophytes close to the ant nests can be therefore seen as an example of indirect myrmecophily.