Allometric allocation in fruit and seed packaging conditions the conflict among selective pressures on seed size

Selective pressures on seed size could vary among the different stages of plant life cycles, so no simple relation could explain a priori its evolution. Here, we determined the relationships between seed size and two fitness components—seed dispersal and survival from predation—in a bird-dispersed tree, Crataegus monogyna. We interpret these relationships in relation to the patterns of mass allocation to fruit and seed components. Selection patterns were assessed at two levels (1) selection pressures on the parent tree; comparing seed dispersal efficiency among individual plants and (2) selection pressures at the individual seed level; comparing seed size variation (i) before and after dispersal, and (ii) before and after postdispersal seed predation. Dispersal efficiency (percentage of seed crop dispersed) was positively correlated with fruit mass and fruit width. Differences in crop size did not offset this effect, and larger seeds were overrepresented in the seed rain relative to the seed pool before dispersal. However, the advantage of larger seeds during the dispersal stage was cancelled later by an opposite selection pressure exerted by seed predators. As a result, smaller seeds had a higher probability of surviving postdispersal seed predation, establishing an evolutionary conflict imposed by the need for dispersal and the danger of being predated. Birds and rodents preferentially selected highly profitable fruits and seeds in terms of the relative proportion of their components. Larger fruits had a higher pulp to seed proportion than smaller ones, and all seeds had the same proportion of coat relative to the embryo-plus-endosperm fraction. Hence, although predator pressures were stronger than disperser ones, larger seeds invested proportionally less in structural defense than in dispersal.     

Selective fruit and seed maturation in Asphodelus albus Miller (Liliaceae)

The fruiting patterns of the rhizomatous perennial Asphodelus albus Miller (Liliaceae) were studied in five populations during 1989 and in one population over 3 years (1988–1990). Fruit/flower (Fr/Fl) ratio and seed/ovule (S/O) ratio varied markedly between populations. Although there were differences between years within the population studied over 3 years, these variations, although statistically signifcant, were less important than those between populations. Neither flowering phenology nor plant size influenced Fr/Fl or S/O ratios. Field experiments tested whether fruit and seed set were pollenor resource-limited. Hand-pollination had no effect on Fr/Fl or S/O ratios, but the reduction of resources by defoliation at the time of flowering decreased both relative components of fecundity. Manipulation of resources by hand-thinning flowers and tiny fruits had no demonstrable effect on these ratios, although brood size of individual fruits was significantly affected. It may be concluded that fruit maturation is resource-limited rather than pollen-limited. Most of the fruits aborted early in the fruiting season, and fruits with higher numbers of developing seeds had a lower probability of abortion than fruits with fewer seeds. Analyses of position effects revealed that the fruits in lower positions in the inflorescence matured preferentially. Furthermore, the two ovules in the same carpel tended either both to fail or both to develop into seeds. The plant's ability to selectively mature only high quality embryos may be limited.     

The influence of ethanol on feeding in the frugivorous yellow-vented bulbul (Pycnonotus xanthopygos)

The sugars in fleshy fruits provide a rich source of energy to frugivorous animals. However, these carbohydrates also serve as a substrate for alcoholic fermentation by yeasts, ethanol being the main by-product of this process. Ethanol ingestion via frugivory thus occurs in a diverse assemblage of invertebrate and vertebrate taxa, including numerous species of birds. We tested the roles of ethanol as an odor cue for resource location by adult yellow-vented bulbuls (Pycnonotus xanthopygos) and as a possible appetite stimulant in feeding trials with artificial food. We hypothesized (1) that the odor of ethanol does not serve as a food-locating cue in diurnal frugivorous passerine birds, and predicted that the choice of food source and the mass of food eaten by such birds will not be influenced by the odor of ethanol. We further hypothesized (2) that food intake in passerine birds is affected by ingestion of ethanol according to its concentration [EtOH], and predicted that food intake will follow a bell-shaped curve in relation to [EtOH]. In accord with hypothesis (1) and its prediction, we found that the odor of ethanol did not affect food preferences, in either ethanol-naïve or ethanol-experienced yellow-vented bulbuls, when presented at concentrations found in naturally ripe fruit (0.0–1.0%); this suggests that the odor of ethanol is not a food-locating cue for the bulbuls. Hypothesis (2) was partially supported, namely at low [EtOH] (0–3%), food intake was constant and at high [EtOH] (3%) food intake decreased, following only the right half of the predicted bell-shaped response. Ethanol-naïve birds showed no preference towards any [EtOH] presented in two-way choice trials. However, daily food intake in ethanol-experienced bulbuls in single option trials decreased by an average of 36% when the artificial food contained the highest tested concentration of ethanol (3.0%). We suggest that decreasing food intake when food ethanol concentration is relatively high may be a means of avoiding intoxication and is related to the ethanol-metabolizing ability of the bird.     

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 diet of Alouatta guariba and Brachyteles arachnoides in Southeastern Brazil: comparison of fruit type, color, and seed size

Fruit is an important food resource for neotropical primates. In this study I compare the fruit diet of sympatric brown howlers (Alouatta guariba) and southern muriquis (Brachyteles arachnoides). Feeding behavior was studied over 12 months and fruit species consumed were identified and assigned to the categories fruit type, fruit color, and seed size. Observed-fruit feeding records were compared with expected records determined from local availability of the fruit of the tree species. I also determined dietary overlap. Fruit consumption occupied 8 and 12% of the feeding time of A. guariba and B. arachnoides, respectively. Fruit from eight tree species were consumed by the former and fruit from twenty-two species by the latter. Patterns of fruit selection of A. guariba and B. arachnoides varied widely. Although howlers and muriquis converge behaviorally by selecting fruit with common attributes (fleshy/unprotected, violet and brown/black-colored), unlike A. guariba, B. arachnoides fed on immature seeds of fleshy/protected and dry fruit. Large seeds were ingested, and defecated intact, by B. arachnoides only. There was little overlap of fruit diet even within categories that had been selected by both, suggesting that dietary divergence is occurring at the interspecific level. Different resource exploitation probably mediates the coexistence of A. guariba and B. arachnoides in low diversity, semideciduous forests, where the environment imposes narrow limits on primate food choices.     

Effect of pollination intensity on fruit and seed set in cacao (Theobroma cacao L.)

We studied the functional relationship between pollination intensity and fruit survival as well as the number of seeds per pod in the tropical tree Theobroma cacao L. on a Forastero Upper-Amazon clone (UPA 409) in Ivory Coast. Cutting the style 24 h after pollination allowed for counting the number of pollen grains deposited on a stigma without affecting fruit set and seed development. Forty-three pollen grains were necessary to reach 50% of maximum fruit set 28 days after pollination. Above 115 pollen grains, the proportion of developing ovaries reached a maximum of 88% 28 days after pollination and 75% at maturity. With fewer than 238 pollen grains per stigma, there was a close relationship between pollination intensity and number of seeds per pod; the pollenseed ratio increased from 1.61 to 3.81 for PI increasing from 30 to 238 pollen grains. For higher pollination intensities, the average number of seeds per pod reached a maximum of 58. The relationship between pollination intensity and seed content was modelled. Results are consistent with the hypothesis that ovules attracted pollen tubes in a similar way regardless of whether or not they had already been reached by another pollen tube.     

白及果实形态与种子结实率相关性研究

该研究以果实大小有明显差异的26个白及果序共87枚果实为材料,分析了果实形态(包括果实重量、长度及直径)与种子结实率的相关性。结果表明:果实重量与结实率的相关性最为密切,相关系数为0.786,达到中等正相关;其次是果实直径,相关系数为0.634;最后是果实长度,相关系数为0.553。对10枚果实及种胚大小测量结果显示,果实形态与种胚大小相关性为弱相关,不同果实种胚大小相差较大,种胚直径和种胚长度平均分别为177.9μm和359.0μm,最大分别为241.9μm和528.6μm,最小分别为64.5μm和114.3μm;单粒白及种子平均重3.90μg,87枚果实中,单枚果实平均种子数量为10 436粒,最大为39 488粒。该研究结果表明果实重量及饱满程度(直径),可以作为评价白及果实质量(种子结实率)的参考指标。     

Satiation of predispersal seed predators: the importance of considering both plant and seed levels

Plants can reduce the fitness costs of granivory by satiating seed predators. The most common satiation mechanism is the production of large crops, which ensures that a proportion of the seeds survive predation. Nevertheless, satiation of small granivores at the seed level may also exist. Larger seeds would satiate more efficiently, enhancing the probability of seed survival after having been attacked. However, a larger seed size could compromise the efficiency of satiation by means of large crops if there were a negative relationship between seed size and the number of seeds produced by an individual plant. We analyze both types of satiation in the interaction between the holm oak Quercus ilex and the chestnut weevil Curculio elephas. Both crop size and acorn size differed strongly in a sample of 32 trees. Larger crop sizes satiated weevils, and higher proportions of the seeds were not attacked as crop size increased. Larger seeds also satiated weevil larvae, as a larger acorn size increased the likelihood of embryo survival. Seedling size was strongly related to acorn size and was reduced by weevil attack, but seedlings coming from large weeviled acorns were still larger. The number and the size of the acorns produced by individual trees were negatively related. Larger proportions of the crop were infested in oaks producing less numerous crops of larger acorns. However, contrary to expectations, these trees did not satiate more effectively at the seed level either. Effective satiation by larger acorns was precluded by larger multi-infestation rates associated to smaller seed crops, in such a way that the proportion of attacked seeds that survived did not vary among trees with different acorn sizes. These results highlight the need of considering satiation by means of large crops and large seeds in studies of predispersal seed predation. Long-term monitoring on individual oaks will help to assess whether there is a trade-off between the number and the size of the acorns and, if it existed, how it could condition the fitness consequences of both types of satiation.     

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.     

Fruit phenology of Prunus jamasakura and the feeding habit of the Asiatic black bear as a seed disperser

We investigated the relationship between the fruit phenology of Prunus jamasakura and the fruit-feeding period of the Asiatic black bear (Ursus thibetanus). The purposes of this study were to determine (1) when bears feed on the fruit of P. jamasakura in relation to phenology; (2) whether ingestion damages seeds; and (3) how ingestion influences seed germination. We assessed the relationship between the phenology of fruit maturation (size, sugar concentration, color, persistence, and germination percentage) and the feeding period of bears in the field, as judged from bear shelves and claw marks. We also compared the germination percentage of seeds ingested by captive bears with that of uningested. Bears fed on the fruit from days 50 to 66 after flowering, when most of the fruits were on the tree and became large, the germination percentage of seeds increased, and the sugar concentration became high. Bears fed only on mature fruits and so obtained high-quality nutrients. Germination tests showed that ingestion of fruits by the bears caused no physical damage to the seeds. Ingested seeds did not show a significant difference in germination percentage from seeds, whose pulp was artificially removed. These results indicate that bears are potentially effective dispersers from a qualitative perspective and, to some extent, from a quantitative perspective.     

The effects of seed weight on growth and competitive ability of Rumex acetosella from two successional old-fields

Summary The effects of seed size on growth, biomass allocation and competitive ability of Rumex acetosella plants grown either individually or in competition were studied in two populations (6 months and 15 years old respectively) sampled from a postcultivation successional gradient. For plants grown individually there were highly significant effects of seed weight on growth after 43 days, with a higher relative growth rate (RGR) observed for plants raised from heavier seeds. However at the end of the experiment, seedlings developed from lighter seeds had a RGR 2 times greater than those from heavier seeds. Final biomass of the two types was not significantly different after 73 days of growth. When plants were grown individually, there were only slight differences between populations, but when grown in monocultures of 4 plants per pot, plants from the old population had higher root and leaf biomass per pot whereas those from the young population had a higher reproductive effort per pot. This suggests that a trade-off between allocation to sexual and vegetative reproduction occurs over successional time. In mixtures of light and heavy seeds, plants from light seeds were shorter, had fewer leaves and lower biomass than plants from heavy seeds, which were also taller and produced more dry matter than plants grown from heavy seeds in monoculture. The significant effects of seed weight and population on biomass parameters persisted unit the end of the experiment. Seedlings from heavy seeds were strong competitors: those from the young population grew better in the presence of neighbors than in monoculture and those from the late successional population suppressed the more the growth of their partners. Seedlings from light seeds were subordinate competitors. These results suggest that seedlings from seeds of different sizes benefit from contrasting ecological conditions and that selection acts on reproductive output along successional gradients.     

Facultative ripening in Hamelia patens (Rubiaceae): effects of fruit removal and rotting

Summary In Costa Rica individual Hamelia patens trees produce fruit throughout the year and experience dramatic changes in rates of fruit removal and rotting. During some moths, most fruits rot because they are not removed. Rotting fruits increase the probability that other fruits on the same infructescence will rot. When removal rates are high, fruits are taken as soon as their seeds become viable but before the fruit is completely ripe. Experimental removal of fruits produced significantly higher ripening rates than on control infructescences. This response allows Hamelia to ripen more fruit and increase the number of fruits taken when dispersers are abundant (e.g., during migration). The proximate mechanism of this response probably includes reallocation of energy conserved when partially ripe fruits are removed. Responding to fluctuating disperser populations likely increases dispersal success and may function as the ultimate cause.     

Seasonal variation in fleshy fruit use and seed dispersal by the Japanese black bear (<Emphasis Type="Italic">Ursus thibetanus japonicus</Emphasis>)

Fruit use by the Japanese black bear (Ursus thibetanus japonicus) and seed clumping in bear scat were studied in central Japan using fecal analyses. Between May and November 2003 and 2004, the life form and fruit size of plants consumed by bears and the species composition and intactness of seeds contained in scat were examined in five transects (approximately 10 km × 10 m) in broad-leaved deciduous forests. In 2003, scats with seeds were found only in the autumn, when fruiting trees and shrubs were abundant. In 2004, scats with seeds occurred intermittently from the summer, when fruiting plants were rare, up to the autumn. Yearly and seasonal variation in fruit use reflects the opportunistic foraging behavior of Japanese black bears. Seven of the nine plant species detected in scats had medium-sized fruits (6–15 mm width), whereas the other two species had relatively large fruits (20–100 mm width). In total, 14,492 seeds were detected, of which 97.6% were intact; the remainder were damaged. Intact seeds of one or two species were found in each scat. The number of intact seeds per scat ranged from 1 to 5476. Japanese black bears seldom digest ingested seeds, thereby contributing to the seed dispersal of their food plants, including species with fruits that are too large to be swallowed by frugivorous birds.     

Interaction between juniper Juniperus communis L. and its fruit pest insects: Pest abundance, fruit characteristics and seed viability

The relationships between the fruit features of Juniperus communis and the presence of fruit pests were studied in Sierra Nevada, SE Spain. The abundance of two insect species — a pulp-sucking scale and a seed-predator wasp — was surveyed with respect both to fruit characteristics and to viability of seeds contained therein. Seed-predator pressure was not significantly related to any fruit characteristics; however, pulp suckers tended to be more abundant in plants with low pulp: seed ratios and high fruit-water content. In addition, fruits with high levels of pulp-sucker attack tended to have higher water content. A multi-factor ANOVA, considering the identity of the plant and the attack of the different pests as factors, showed that plant identity accounts for most of the variation in fruit characteristics. The viability of seeds tended to be lower in plants strongly attacked by both pests. Fruits attacked by seed predators showed significantly lower proportions of viable and unviable seeds than did unattacked fruits. Seed viability was also lower in those fruits heavily attacked by pulp suckers, but this pattern is strongly mediated by plant identity. Pest activity proved to be clearly associated with a direct decrease in juniper reproductive capacity. This loss involved a reduction of the viable-seed number, mainly related to the seed predator, as well as a reduction of fruit attractiveness to frugivorous dispersers, related to the pulp sucker.     

A field observation on color selection by New World sympatric primates,Pithecia pithecia and Alouatta seniculus

This work characterizes differences in selection ofTalisia retusa fruits by two sympatric Neotropical primates,Pithecia pithecia (white-faced sakis) andAlouatta seniculus (red howlers). Color appears to be the criterion by which fruits were selected. Greenish fruits were mainly eaten byPithecia, while yellowish fruits byAlouatta. The characteristics of these primates in relation to seed predation and seed dispersal are discussed in the context of theTalisia retusa fruit color spectrum. Furthermore, a possible differential acquisition of chemical components, like tannins, is hypothetically treated considering the variation in fruit color.     

Correlated evolution of fig size and color supports the dispersal syndromes hypothesis

The influence of seed dispersers on the evolution of fruit traits remains controversial, largely because most studies have failed to account for phylogeny and or have focused on conservative taxonomic levels. Under the hypothesis that fruit traits have evolved in response to different sets of selective pressures by disparate types of seed dispersers (the dispersal syndromes hypothesis), we test for two dispersal syndromes, defined as groups of fruit traits that appear together more often than expected by chance. (1) Bird syndrome fruits are brightly colored and small, because birds have acute color vision, and commonly swallow fruits whole. (2) Mammal syndrome fruits are dull-colored and larger on average than bird syndrome fruits, because mammals do not rely heavily on visual cues for finding fruits, and can eat fruits piecemeal. If, instead, phylogenetic inertia determines the co-occurrence of fruit size and color, we will observe that specific combinations of size and color evolved in a small number of ancestral species. We performed a comparative analysis of fruit traits for 64 species of Ficus (Moraceae), based on a phylogeny we constructed using nuclear ribosomal DNA. Using a concentrated changes test and assuming fruit color is an independent variable, we found that small-sized fruits evolve on branches with red and purple figs, as predicted by the dispersal syndromes hypothesis. When using diameter as the independent variable, results vary with the combination of algorithms used, which is discussed in detail. A likelihood ratio test confirms the pattern found with the concentrated changes test using color as the independent variable. These results support the dispersal syndromes hypothesis.     

The fate of flowers and fruits of Cornus sanguinea L. in three contrasting Mediterranean habitats

Summary In the Mediterranean region of southern France, the fate of flowers and fruits of Cornus sanguinea, a deciduous shrub, was studied in three contrasting habitats: (1) an abandoned olive grove, (2) the edge and (3) the interior of a deciduous forest. Abundance of flowering and fruiting of C. sanguinea differed widely between the three habitats. The fate of the flowers, on the other hand, did not differ significantly between habitats. Taking all three habitats together, 77% of the flowers aborted, 23% initiated fruits, 6% developed mature fruits, and 3% produced ripe fruits that were eaten by birds. Likewise, there were for the most part no significant differences in the fate of immature and mature fruits between the habitats. On average, 58% of the immature fruits were abscised undamaged, 24% matured, 10% were damaged by insects and 8% dried up. Of the mature fruits, on average, 51% were eaten by birds, 23% damaged by insects, 20% dried up and 6% fell to the ground undamaged. Independent of habitat conditions, C. sanguinea seems to regulate the quantity of its seed crop primarily by limiting the number of flowers and secondarily by aborting surplus immature fruits, and the number of flowers is mainly controled by resource availability and genetic factors. In the case of C. sanguinea, both the fruit/flower ratio and the proportion of flowers producing mature fruits that are eaten by birds remain constant over a wide range of environmental conditions.     

Effect of the synthetic auxin 2,4-DP on fruit development of loquat

Application of the butylglycol ester of 2,4-DP increased final fruit size in loquat without causing fruit thinning. Its effectiveness depended on the concentration applied and treatment date. When applied at the onset of the cell enlargement fruit stage, 25 mg l^–1 was the most effective treatment. Fruit diameter distribution showed a significant shift to the larger size for treated trees; further, fruit colour break and maturation were encouraged and harvest time was earlier than in untreated trees.     

Effects of coppicing on demographic structure, fruit and seed set in Orchis mascula

Opening of the forest canopy may result in higher fitness of understory plants due to increased light intensity on the forest soil and increased nutrient availability. Especially, tuberous orchids may profit from increased light as photosynthetic assimilation products and nutrients are accumulated in their belowground storage organs. We investigated the effects of coppicing on demographic structure and fruit and seed set in 15 populations of the tuberous, perennial orchid Orchis mascula, seven of which were located in undisturbed, shaded woodland and eight in recently coppiced woodland. Coppicing resulted in a massive increase in flowering and in increased fruit set. On average, 42.9% of all individuals consisted of flowering plants in coppiced woodland, whereas in undisturbed woodland only 20.8% of all individuals flowered. The percentage fruit set varied between 20% and 55% in coppiced woodland and was strongly related to population size (measured as the number of flowering individuals). Mean fruit weight and fruit size were also significantly higher in populations located in coppiced woodland than in undisturbed woodland, whereas the proportion of viable seeds was not related to woodland type. Given that O. mascula is dependent on seed set to fill space with new individuals, these results suggest that the long-term survival of this species strongly depends on frequent opening of the canopy.     

Effect of ovular receptivity on seed set and fruit development in cucumber (Cucumis sativus L.)

Summary Seed set and fruit development in cucumber (Cucumis sativus L.) were studied in relation to female flower receptivity from day — 2 before anthesis to day + 2 after anthesis. The female cucumber flower is protogynous. The pistil was receptive 2 days before anthesis. The iso-electric focusing (IEF) patterns of the stigma/style proteins, were identical from day -5 to day +2. In pollinated flowers in vivo germination and pollen-tube growth in the ovary were affected by pistil age from day -2 to day +2. In addition, differences in sectorial filling in full seeds were observed within the fruits. A negative correlation was observed between the frequency of fertilized ovules in the pedoncular part of the fruit and ovary length at the time of pollination. In the whole fruit, significant differences in the number of full seeds and fruit size at maturity were found, and these were observed to be correlated with the various stages of female flower maturation at pollination. The day -2 and day +2 stages yielded the smallest fruits with few full seeds compared to the day -1, day 0 and day +1 stages, which had the biggest fruits and a large number of full seeds. A strong positive correlation was found between total seed number (including full and empty seeds), fruit length and weight at maturity. All these results suggest that both seed set in the different parts of the fruit and fruit development are controlled by ovular receptivity rather than by stigma/style receptivity.