DEVELOPMENTAL AND GENETIC SOURCES OF SEED WEIGHT VARIATION IN RAPHANUS RAPHANISTRUM L. (BRASSICACEAE)

Seed weight is known to have a marked impact on emergence and post-emergence productivity in wild radish (Raphanus raphanistrum). In this paper, I describe several levels of seed weight variation in plants taken from a natural population in Hamden, Connecticut. Six maternal plants from the 1981 season were analyzed in detail: the weights and positions of all seeds within a fruit were recorded, and some of these seeds were used the following summer for competition studies and progeny analysis. Within a plant, average seed weight decreased as the number of seeds within a fruit increased, suggesting that developing embryos compete for maternal resources. Seed weight also varied significantly among the six maternal plants used in the study. Comparison of the average weights of seeds produced by offspring of those six plants with the average weights of seeds borne by the maternal plant revealed a significant genetic component to seed weight variation. Seed weight varied up to six-fold within single fruits of R. raphanistrum; large seeds tend to occur near the pedicel or in the middle positions. Seed size variation seen within single fruits is of sufficient magnitude to result in differential reproductive output among closely related seeds under competitive field conditions.     

VARIATION IN AVERAGE SEED SIZE AND FRUIT SEEDINESS IN A FRUIT CROP OF A GUANACASTE TREE (LEGUMINOSAE: ENTEROLOBIUM CYCLOCARPUM)

A detailed examination of the ordinary 1980 fruit crop of an ordinary large adult guanacaste tree (Enterolobium cyclocarpum) in Guanacaste Province, Costa Rica (1980) shows that for fruits with about 6-16 seeds (range, 1-18 seeds per fruit) there is about ***1 g dry weight fruit tissue per seed (seeds averaging about 820 m each). Fruits that have about 1-5 seeds on average have 1.5-3 times as much dry fruit tissue per seed as do the seed-rich fruits, and also contain seeds that weigh about 8% more on average than the seeds in the seed-rich fruits. The average seed weight per fruit was found to decrease slightly if all sizes of fruits are considered, but to remain essentially constant over the range of 7-16-seeded fruits. This category contained 78% of the fruits. These findings suggest that different parts of the seed crop may end up in different dispersers and dispersers with different preferences for fruit seediness and tolerances for seed size may remove different portions of the seed crop.     

REPRODUCTIVE CAPACITY AND SEED SIZE IN LUPINUS TEXENSIS

The average number of ovules produced per individual of Lupinus texensis is much greater than the average number of seeds per plant. Each plant produces approximately 2,000 ovules but only 2.5% develop into seeds. One fourth of the seeds is lost due to abortion and 0.3% is lost due to predation on the plant. Mature seeds from this population exhibit a five-fold range in weight, from 10 to 56 mg. The distribution of seed weights in the field population is skewed and leptokurtic. Seed wt is positively correlated with both seed germination and seedling survivorship. Heritability of seed wt is 0.09. There is no correlation between average seed wt per plant and total number of seeds per plant, seeds per pod, or legumes per plant.     

Seed weight and germination behavior of the submerged plant Potamogeton pectinatus in the arid zone of northwest China

Variation in seed weight is common within and among plant species, but few studies have attempted to document the pattern of seed weight and germination attributes for aquatic macrophytes at a large scale. This study examined within‐species variation in seed weight and germination attributes and the effects of environmental factors on seed traits of the submerged plant Potamogeton pectinatus in the arid zone of northwest China. Our results showed that the average seed weight was 0.24 g per 100 seeds with a coefficient of variation (CV) of 28.4% among the eight P. pectinatus populations. The total germination fraction of seeds of P. pectinatus was relatively poor, less than 35% in seven P. pectinatus populations, and the lowest germination percentage found was only 2%. There were significant differences in seed weight, time to onset of germination, and total germination fraction among the eight different populations. Hierarchical partitioning analysis showed a strongly positive correlation between seed weight and water temperature and pH. Seed weight and the maternal environmental factors significantly affected both time to initiation of germination and total germination fraction. Our results suggest that (1) seed weight variation in P. pectinatus primarily is the result of temperature variation during fruit development; (2) relatively poor germination fraction suggests that seeds are relatively unimportant in the short‐term survival of populations and that it may be another adaptive trait allowing plants to take place in the right place and at the right time, especially in harsh environment; and (3) variation in seed germination traits should be determined by local environmental and intrinsic factors that interact in a complex fashion.     

Seed size selection by olive baboons

Seed size is an important plant fitness trait that can influence several steps between fruiting and the establishment of a plant’s offspring. Seed size varies considerably within many plant species, yet the relevance of the trait for intra-specific fruit choice by primates has received little attention. Primates may select certain seed sizes within a species for a number of reasons, e.g. to decrease indigestible seed load or increase pulp intake per fruit. Olive baboons (Papio anubis, Cercopithecidae) are known to select seed size in unripe and mature pods of Parkia biglobosa (Mimosaceae) differentially, so that pods with small seeds, and an intermediate seed number, contribute most to dispersal by baboons. We tested whether olive baboons likewise select for smaller ripe seeds within each of nine additional fruit species whose fruit pulp baboons commonly consume, and for larger seeds in one species in which baboons feed on the seeds. Species differed in fruit type and seed number per fruit. For five of these species, baboons dispersed seeds that were significantly smaller than seeds extracted manually from randomly collected fresh fruits. In contrast, for three species, baboons swallowed seeds that were significantly longer and/or wider than seeds from fresh fruits. In two species, sizes of ingested seeds and seeds from fresh fruits did not differ significantly. Baboons frequently spat out seeds of Drypetes floribunda (Euphorbiaceae) but not those of other plant species having seeds of equal size. Oral processing of D. floribunda seeds depended on seed size: seeds that were spat out were significantly larger and swallowed seeds smaller, than seeds from randomly collected fresh fruits. We argue that seed size selection in baboons is influenced, among other traits, by the amount of pulp rewarded per fruit relative to seed load, which is likely to vary with fruit and seed shape.     

Seed Type and Seed Size Variation in the Heteromorphic Saltmarsh Annual Spergularia salina along the Coast of Sweden

Abstract: We investigated habitat characteristics, seed type and seed size variation between and within 41 populations of the heteromorphic saltmarsh annual Spergularia salina. 95% of the investigated populations were truly heteromorphic regarding unwinged or winged seeds but either of the seed types could dominate in individual populations. 39% of all seed capsules contained mixed seed types, in 43% of capsules the seeds were exclusively unwinged, and in 18% only winged seeds were found. In a closer survey in one of the populations (at Tullgarn, 5850'N, 1737'E), albeit predominantly heteromorphic, again capsules either contained mostly unwinged or winged seeds: 59% of the capsules had mixed contents, 21% contained exclusively unwinged seeds, and 21 % produced only winged seeds. Seed weight varied up to 10-fold, with an overall mean of 77.2μg. 27% of this variation was due to variation among populations, 52% was explained by variation between individuals, and 21% was due to variation within individuals. Seed weight was positively correlated with the frequency of winged seeds per capsule, especially within populations, and negatively correlated to seed number. The frequency of winged seeds in each capsule and seed weight was positively correlated with population density both between and within populations.     

Effects of seed size on seedling size in Virola surinamensis; a within and between tree analysis

Summary We conducted a greenhouse study of the effects of initial seed mass on seedling characteristics in a Panamanian population of Virola surinamensis, a canopy tree in which mean seed mass of different individuals ranges from 1.34 to 4.04g. The system is of particular interest because birds preferentially eat fruits of small-seeded plants, leaving seedlings of large-seeded individuals under conditions of potentially severe sibling competition (Howe and Vande Kerckhove 1980).Effects of differences of mean seed mass between trees are explored using an analysis of variance, while effects of seed-mass variation within crops are demonstrated with a regression analysis. A two-way analysis of variance decisively shows effects of parental source and light condition on seedling height, leaf length, and dry shoot mass (all P<0.0001). A posteriori tests show that differences in seedling characteristics reflect differences in initial seed mass, with especially strong differences apparent in shoot mass. Regression of seedling characteristics on initial seed mass shows that variation of seed size within a crop is sufficient to influence shoot mass at 15 weeks (P<0.0001).Effects of size differences of seeds that land adjacent to each other, either under the parent or in monkey droppings, are documented with growth of pairs of seedlings in pots. Differences in shoot height and mass at 15 weeks are evident when seeds of average size differ by only 0.2 g, and dramatic differences are evident when paired seeds differ by an average of 1.5 g. Seedlings grow more when isolated than when planted with conspecifics.These experimental results offer indirect support for the hypothesis that small-seeded Virola parents secure an advantage in reproduction through differential dispersal, while large-seeded plants produce more competitive seedlings under their own crowns — an advantage most likely to be of importance when frugivores are scarce.     

濒危植物梵净山冷杉的球果与种子性状研究北大核心CSCD

梵净山冷杉(Abies fanjingshanensis)为国家一级保护濒危植物。为了揭示梵净山冷杉球果发育和成熟过程的性状特征,以确定成熟球果的适宜采种期。该研究对梵净山冷杉自然生长区内成年结实母树的球果进行跟踪观测和定期采集,通过物理解剖和形态学参数测定,比较分析不同采种期梵净山冷杉的球果、种鳞和种子的性状差异。结果显示:(1)梵净山冷杉的球果每年7月中旬开始形成,10月中旬开始成熟,发育早期至发育后期球果的长度和宽度显著增加,发育后期至成熟期形态和颜色均无明显变化。(2)成熟球果平均长、宽、鲜重、干重和相对含水量分别为7.18 cm、3.84 cm、36.98 g、20.33 g和45.06%。(3)成熟球果的平均种鳞层数为30.76层,种鳞总数为250.67片,平均出种量436.67粒,平均种子饱满率82.49%。(4)成熟球果基部、中部、上部的种子性状不同,且球果中部的种子性状参数最大,饱满率最高,种子平均长、宽、厚分别为9.14、2.30、2.37 mm,千粒重11.44 g。研究表明,梵净山冷杉的球果从形成至成熟过程约3个月,成熟球果的最佳采种时间为10月下旬,发育后期和成熟球果的形态和颜色差异不明显,不能以颜色和形态作为球果成熟的判定依据;种鳞与球果的发育和成熟同步,球果中部的种鳞能够完全发育,且形状参数最大,孕育的种子最饱满;梵净山冷杉成熟球果出种量大、饱满率高,但是种子小、重量轻。     

Parsnip (Pastinaca sativa) seed production: effects of seed crop plant density, seed position on the mother plant, harvest date and method, and seed grading on embryo and seed size and seedling performance

Parsnip seeds from four years of seed production (1980–83) were used to examine the effects of position of the seed on the parent plant and several cultural factors on seed and embryo growth and seedling performance. Maximum seed dry weight was reached c. 50 days after flowering (DAF) but embryos continued to grow for a further 7–15 days. The coefficient of variation (C.V.) of seed weight fell from c. 35–40% at 40 DAF, to about 20–25% at maturity. The corresponding figures for C.V. of embryo length were 25–35% at the early harvest and 12–15% at maturity. The effects of seed position, plant density, harvest method, seed grading after harvest on seedling emergence and weight characters were small. Harvesting seeds c. 60 DAF gave, on average, 3.3 days earlier emergence, c. 3.6 days lower ‘spread’ of emergence time and c. 9% higher seedling emergence than harvesting 2 wk earlier. Late harvesting of the seed crop reduced the C.V. of seedling weight from 53 to 46%, on average, but there were variations in response from year to year. Seedling emergence and seedling weight characters were more closely related to embryo length than to seed weight characters.     

Quantitative genetic analysis of embryo heterosis in faba bean (Vicia faba L.)

Seeds, i.e. embryos, may be genetically different from either of their parents and moreover may express their own heterosis. The objective was to genetically analyse embryo heterosis for their own weight (i.e. seed weight) in comparison with their seedlings’ heterosis, taking the large-seeded crop (Vicia faba L.) as model. A specific diallel mating scheme was used, based on four parental lines, creating 76 seed genotypes in generations P, F₁, F₂ and BC. Mature seed weight was assessed for these embryo genotypes in 3 years at one German location, and young plant biomass yield of seedlings emerging from these seeds in two greenhouse experiments. The quantitative genetic analysis showed an average of 10.6% mid-parent heterosis for mature seed weight and 14.5% mid-parent heterosis for juvenile biomass. In both traits, the embryos contributed markedly and significantly via their own genes to the genetic variation. For mature embryo weight heterosis, apparently the parental difference in seed weight was decisive, whereas for juvenile biomass heterosis, genetic unrelatedness of parents had priority.     

FACTORS INFLUENCING SEED WEIGHT IN IMPATIENS CAPENSIS (BALSAMINACEAE)

I analyzed within- and between-plant variation in seed weight from one greenhouse and four natural populations of Impatiens capensis Meerb. Most variation in seed size in natural populations occurred within populations and within plants. Weights of seeds from cleistogamous flowers on greenhouse plants increased with height on the plant and proximity to the main stem, a result that may be related to the local supply of photosynthate. Mean seed weight in the natural populations declined during seed set, then increased again. Seeds from chasmogamous flowers were significantly larger in one of the four populations. Maternal parents differed significantly in both mean seed weight and the relative size of the two types of seeds. Combined genetic and general environmental effects accounted for 26% of the total phenotypic variance, which provides an outside estimate of the heritability of this character.     

Inflorescence characteristics,seed composition,and allometric relationships predicting seed yields in the biomass crop Cynara cardunculus

Cynara (Cynara cardunculus) is a perennial C₃ herb that has its potential as bioenergy crop. This paper aims (a) to derive empirical relationships to predict cynara seed yield per head and per unit area, avoiding laborious extraction of seeds from the complex structure of its inflorescences; (b) to determine the head‐weight distribution per unit area, the seed composition and the oil profile of cynara seeds; and (c) to estimate the range of cynara biomass, seed and oil yield in representative parts of Greece. We analyzed 16 field experiments, varying in crop age and environmental conditions in Greece. Seed yield per head (SY_head) can be accurately predicted as a linear function of dry head weight (H_w): SY_head=0.429·H_w?2.9 (r²=0.96; n=617). Based on this relationship, we developed a simple two‐parameter equation to predict seed yield per unit area (SY): SY=HN·(0.429·μ?2.9), where μ is the mean head weight (g head^?1) and HN is the total number of heads per unit area, respectively. The models were tested against current and published data (n=180 for head‐level; n=35 for unit area‐level models), and proved to be valid under diverse management and environmental conditions. Attainable cynara seed yields ranged from 190 to 480 g m^?2 yr^?1, on dry soils and on aquic soils (shallow ground water level). This variation in seed yield was sufficiently explained by the analyses of head‐weight distribution per unit area (small, medium and large heads) and variability of seed/head weight ratio at head level. Seed oil concentration (average: 23%) and crude protein concentration (average: 18.7%) were rather invariant across different seed sizes (range: 26–56 mg seed^?1) and growing environments.     

Resource distribution and the trade-off between seed number and seed weight: a comparison across crop species

In grain crops, total sink capacity is usually analysed in terms of two components, seed number and individual seed weight. Seed number and potential individual seed weight are established at a similar timing, around the flowering period, and seed weight at maturity is highly correlated with the potential established earlier. It is known that, within a species, available resources during the seed set period are distributed between both yield components, resulting in a trade-off between seed number and seed weight. Here we tested if this concept could apply for interspecific comparisons, where combinations of numbers and size across species could be related to the total available resources being either allocated to more seed or larger potential individual seed weight during the seed set period. Based on this, species differences in seed weight should be related to resource availability per seed around the period when seed number is determined. Resource availability per seed was estimated as the rate of increase in aboveground biomass per seed around the period of seed set. Data from 15 crop species differing in plant growth, seed number, seed weight and seed composition were analysed from available literature. Because species differed in seed composition, seed weight was analysed following an energy requirement approach. There was an interspecific trade-off relationship between seed number per unit of land area and seed weight (r = 0.92; F(1, 13) = 32.9; n = 15; P < 0.001). Seed weight of different species was positively correlated (r = 0.90; F(1, 13) = 52.9; n = 15; P < 0.001) with resource availability per seed around the seed set period. This correlation included contrasting species like quinoa (Chenopodium quinoa; ∼100000 seeds m⁻², ∼4 mg equivalent-glucose seed⁻¹) or peanut (Arachis hypogaea; ∼800 seeds m⁻², ∼1000 mg equivalent-glucose seed⁻¹). Seed number and individual seed weight combinations across species were related and could be explained considering resource availability when plants are adjusting their seed number to the growth environment and seeds are establishing their storage capacity. Available resources around the seed set period are proportionally allocated to produce either many small seeds or few larger seeds depending on the particular species.     

POLLEN AND OVULE SOURCES AFFECT SEED PRODUCTION OF LOBELIA CARDINALIS (LOBELIACE AE)

To examine the breeding system and components of male and female reproductive success in the hermaphroditic plant Lobelia cardinalis, we performed three crossing experiments with plants taken from natural populations. The experiments were designed to determine if the crossing success of plants as pollen and ovule parents was affected by the distance among mates, including self-pollinations and pollinations between populations; to determine if plants differed in their abilities to sire or mature seed; and to determine if there was a correlation between a plant's success at siring and maturing seed. Selfpollinations resulted in significantly fewer seeds per fruit and significantly smaller seeds. There were no significant differences in germinability between selfed and outcrossed seeds. Distance among parents within a population did not affect any of the traits. Outcrosses within and between population produced similar numbers of seeds per fruit, similar seed weights, and similar germination success. There were highly significant differences among maternal plants in all three experiments in the number of seeds they matured, mean seed weight, and seed germinability. The maternal parent was the most important factor determining seed production, but there were also significant differences among paternal plants in the number of seeds they sired (all three experiments), in the germinability of the seeds they sired (two experiments), and in the sizes of seeds they sired (one experiment). Our results indicate that differences in success of Lobelia plants as male parents cannot be due solely to their relatedness to the female parent.     

Discrimination among pinyon pine trees by Clark's Nutcrackers: effects of cone crop size and cone characters

Summary The influences of Colorado pinyon pine (Pinus edulis) cone crop size, cone and seed weight, cone length, number of seeds per cone, number of viable seeds, and percent viable seeds on the foraging behavior of avian seed dispersal agents were examined in field and laboratory settings. In the field, there was a significant positive relationship between cone number per tree and both the absolute number of cones and the percentage of the cone crop from which seeds were harvested. Cone weight and the number of viable seeds were also significantly related to seed harvest intensity. Laboratory experiments examined the relationship between crop size and cone characters on seed harvest by 18 Clark's Nutcrackers (Nucifraga columbiana). Nutcrackers were offered a choice of two tree types: one with 20 cones attached, and another with 10 cones attached. Significantly more birds chose to remove seeds first from the tree with 20 cones than the tree with 10 cones. In timed trials, they also harvested seeds from significantly more cones on the tree with the higher cone density. In the laboratory, cones chosen for seed removal by the nutcrackers had significantly more viable seeds, more seeds, and were longer compared to cones that were not chosen. Such discriminatory foraging behavior may increase avian foraging efficiency and result in differential reproductive success of pinyon pines. This behavior may therefore influence the evolution of pinyon pine reproductive traits.     

Structure of Protochlorophyll-containing Plastids in the Inner Seed Coat of Pumpkin Seeds (Cucurbita pepo)

The inner seed coat of seeds of Cucurbita pepo L. cv. Ohlsens Enke Köks was used to study the development of protochlorophyll-containing plastids with an abnormal ultrastructural composition. The pumpkins were harvested at different stages during fruit development and they thus contained seeds with different developmental stages. The dry weight of seeds of the developmental stages used varied from 0.04 g to 0.3 g. Such a series of seeds with decreasing water content indicating increasing maturity contained different amounts of protochlorophyll, from 0.20 μg/g fresh weight to 500 μg/g fresh weight. The ultrastructure of the protochlorophyll containing plastids changed greatly during development. In young seeds with a low content of protochlorophyll, regular prolamellar bodies were found and starch grains filled most of the plastids. During development the starch content decreased and the prolamellar bodies increased in size and lost their regularity. During maturation the plastids accumulated plastoglobuli, probably containing protochlorophyll, and finally the internal structure of the prolamellar body tubular complex was lost.     

Seed size variation and its effect on germination and seedling performance in the clonal herb Convallaria majalis

Variation in seed size is common both within and among plant species. This study examined within-species variation in seed weight, and its implications for some components of fitness in the clonal herb Convallaria majalis. This species produces berries containing 4.1 seeds on average. The average seed weight was 16.5 mg, with a coefficient of variation of 32.7%. Seed packaging in fruits was on average 12.5%, and showed a slight tendency to increase with fruit weight. A trade-off was found between seed weight and seed number both within fruits and within ramets. The probability and timing of germination was not influenced by seed size. A field experiment and indirect evidence suggested that post-dispersal seed predation was not related to seed size. Increasing seed weight conferred an advantage to developing seedlings. This advantage was enhanced if a seedling was growing in the close vicinity of a seedling of another species. It is suggested that seed size variation in C. majalis primarily is the result of resource variation during fruit development. A conflict between parents and offspring may however contribute to increase seed size variation.     

The genetics and ecology of seed size variation in a biennial plant,Hydrophyllum appendiculatum (Hydrophyllaceae)

The goal of this study was to elucidate the sources of seed size variation in Hydrophyllum appendiculatum, an outcrossing, biennial plant. The genetic basis of seed size variation was examined with a diallel breeding design. The analysis did not reveal any evidence for additive genetic variance, suggesting that seed size could not evolve in response to natural selection. A series of greenhouse experiments was conducted to determine the sensitivity of seed weight to a number of ecological variables. Seed weight was affected by inbreeding depression: seeds produced by self-pollinations were significantly lighter that outcrossed seeds. Maternal plants did not differentially provision seeds that were the result of crosses between subpopulations (separated by 300 m) or between populations (separated by 1.7 km). Mean seed size was independent of the number of outcrossed pollen donors (one vs. many) that sired seeds on an inflorescence; however, the variance was greater on inflorescences pollinated by multiple donors. Direct manipulations of the abiotic environment showed that seed size was greater on plants growing under full sunlight compared to shaded plants. Seed size was unaffected by soil type, fertilizer addition, or defoliation. Finally, I determined the effect of varying pollination intensity at the level of a single inflorescence, and at the whole plant level. Seed weight was greatest on plants that had only 1 and 5 inflorescences pollinated, and least on those that had 10 and 20 pollinated. At the inflorescence level, seed weights were greatest on those where all flowers were pollinated, compared to inflorescences where only half of the flowers were pollinated. Perhaps the greatest contributor to variance in seed size in this species was the temporal decline within plants through the flowering season. These results indicate that maternal plants are not capable of producing uniform seed crops. Rather, the final distribution of seed size produced by plants within a population will necessarily vary and be the result of pollination effects, heterogeneity in the abiotic environment, and developmental constraints.     

Self- and open-pollination as factors influencing seed quality in leek (Allium porrum)

The effect of self-pollination compared with open-pollination on leek seed weight, and subsequent seedling emergence and growth were observed for two years of seed production. Open-pollination gave higher mean seed weights and lower seedling losses during plant establishment than self-pollination. Depending upon the year, the individual plant and pollination method, between 2. 4 and 56.6% of the variation in seedling weight c. 4–5 wk after sowing under controlled conditions was accounted for by differences in seed weight and time of seedling emergence. In 1984 but not 1983, a greater proportion of the variation in seedling weight was accounted for by seed weight and time of seedling emergence in open than self-pollinated seeds.     

Fruit production in Sorbus aucuparia L. (Rosaceae) and pre-dispersal seed predation by the apple fruit moth (Argyresthia conjugella Zell.)

Variation in fruit production and pre-dispersal seed predation by Argyresthia conjugella was studied in␣four populations of Sorbus aucuparia in northern Sweden.␣The number of infructescences, fruits per infructescence, consumed seeds and developed unattacked seeds per fruit were scored in marked trees from 1984 to 1990. The results showed that the number of fruits produced in each population determined the number of seed predators occurring in the host population, as the yearly number of seed predators was significantly and positively correlated with yearly number of fruits, in all but one population. The seed predators showed a delay in response to variation in number of fruits produced. This lag in response resulted in a large proportion of fruits being attacked and seeds consumed in a bad fruiting year that followed a good fruiting year, and vice versa. The proportion of fruits attacked and seeds consumed was largest in the population showing the greatest between-year variation in fruit production and lowest in the population showing the lowest between-year variation in fruit production. Furthermore, the individuals within the former population were synchronised, while they were not in the latter population. These results contradict one of the possible explanations of mast-seeding, where large synchronised between-year variation is supposed to reduce pre-dispersal seed predation. Instead, differences in attraction of the seed predator to differences in fruit crop size could explain the observed difference in seed predation between the two populations with opposite fruiting patterns. Within each population, irrespective of year, the proportion of fruits attacked and seeds consumed was independent of a tree's fruiting display. Therefore, trees with high fruit production, despite harbouring the largest number of seed predators, produced the largest number of developed seeds in absolute numbers, compared to trees that produced few fruits. Received: 25 February 1996 / Accepted: 30 November 1996