EFFECT OF DENSITY ON SECONDARY SEX CHARACTERISTICS AND SEX RATIO IN SILENE ALBA (CARYOPHYLLACEAE)

A field survey of plant and flower sex ratio and secondary sex characteristics was made in Silene alba. Female-biased plant sex ratios were found, as seems typical for the species. Sex ratio distribution correlated with a gradient of soil moisture (with the more moist area having a more female-biased ratio) and with changes in the density of Silene (intermediate and higher density areas having greater female bias). The floral sex ratio was significantly female-biased only at the site that was most female-biased in terms of plant sex ratio. Otherwise the population of flowers was significantly male-biased. Male and female plants harvested from the field differed in secondary sexual characteristics. Males had more flowers and invested proportionately more biomass in leaf, but less in root, stem and reproductive tissue than did females. Although both males and females were larger in terms of total dry weight at the moist site, males produced more flowers at the driest (high density) site. Here the female bias in plant sex ratio was intermediate, but the floral sex ratio was significantly male-biased. A glasshouse experiment was performed in which plants were grown at four densities. Density significantly influenced plant survivorship and the probability of flowering, and increased female bias in the pots, but it did not affect patterns of biomass allocation in flowering plants. Patterns of male and female biomass allocation did not differ in the experiment, except in terms of reproductive allocation (greater in females) and allocation to leaf, greater in males, but only at the lowest density. This work urges caution in interpreting differences between males and females in the field as secondary sex characteristics, since we find such properties to be overlapping under experimental conditions. It supports the idea that males and females of a species may sustain different reproductive output under differing conditions.     

GENOTYPIC AND ENVIRONMENTAL COMPONENTS OF VARIATION IN GROWTH AND REPRODUCTION OF FISH HEMICLONES (POECILIOPSIS: POECILIIDAE)

The frozen-niche-variation model was proposed to account for the coexistence of genetically related clones in naturally occurring unisexual populations. This model is based on two assumptions: 1) ecologically different clones have multiple independent origins from sexual ancestors; and 2) the population of sexual ancestors contains genetic variability for ecologically relevant traits. To test these assumptions, we produced 14 new “hemiclones” (nonrecombining haploid genotypes) of fish (Poeciliopsis: Poeciliidae). Our ability to synthesize many new hemiclones demonstrates the feasibility of multiple independent origins of nonrecombining genotypes. A substantial proportion (10–50%) of the phenotypic variation among hemiclones in size at birth, juvenile growth rate, and fecundity had a genetic basis. Thus, we conclude that multiple origins can give rise to an assemblage of genetically distinct hemiclones, each with a unique combination of life-history traits. Additionally, a comparative analysis of two natural hemiclones revealed that the synthetic strains represent a broad field of variation from which natural hemiclones can be selected.     

SEX RATIO SELECTION AND THE EVOLUTION OF ENVIRONMENTAL SEX DETERMINATION IN LABORATORY POPULATIONS OF MENIDIA MENIDIA

What happens when a population with environmental sex determination (ESD) experiences a change to an extreme environment that causes a highly unbalanced sex ratio? Theory predicts that frequency-dependent selection would increase the proportion of the minority sex and decrease the level of ESD in subsequent generations. We empirically modeled this process by maintaining five laboratory populations of a fish with temperature-dependent sex determination (the Atlantic silverside, Menidia menidia) in extreme constant temperature environments that caused highly skewed sex ratios to occur initially. Increases in the minority sex consistently occurred from one generation to the next across all five populations, first establishing and then maintaining a balanced sex ratio until termination of the experiment at 8 to 10 generations. The extent to which the level of ESD changed as balanced sex ratios evolved, however, was not consistent. Two populations that experienced high temperatures each generation displayed a loss of ESD, and in one of these ESD was virtually eliminated. This suggests that temperature-insensitive, sex-determining genes were being selected. In populations maintained in low temperature environments, however, the level of ESD did not decline. Instead, the response of sex ratio to temperature was adjusted upward or downward, perhaps by selection of sex-determining genes sensitive to higher (or lower) temperatures. The two different outcomes at low versus high temperatures occurred independent of the geographic origin of the founding population. Our results demonstrate that ESD is capable of evolving in response to selection.     

EFFECTS OF DISTANCE BETWEEN POLLEN DONOR AND POLLEN RECIPIENT ON FITNESS COMPONENTS IN ESPELETIA SCHULTZII

The effect of flowering time and of distance between pollen donor and pollen recipient on the proportion of filled achenes, aborted seeds, and seedling survivorship was studied in populations of Espeletia schultzii in the Venezuelan Andes. Hand-pollinations were performed in two different years and at different times during the flowering season. Pollinations within-population included crosses between plants within a few meters to a maximum of 500 m apart and between-population pollinations included crosses between plants more than 10 km and up to 78 km away. Late in the flowering season, plants suffered a reduction in the proportion of filled achenes and an increase in aborted seeds. The proportion of filled achenes and aborted seeds did not vary significantly between the distances between pollen and ovules in crosses done early in the flowering season. However, the proportion of filled achenes in crosses between individuals within 30 m distance were very variable, either very successful or not successful. Beyond 30 m, for some of the populations, the frequency of unsuccessful crosses was almost 0. This pattern was observed at three elevation sites in two consecutive years. When crosses were done late in the flowering season, the proportion of filled achenes was lower in crosses between close individuals and between very distant ones. Seedling survivorship varied significantly between the crossing distances at the higher elevation only. The results show that a clear effect of the incompatibility alleles on the crossing success of the plants is present in the different populations examined, while they suggest that the effect of inbreeding or outcrossing depression is less clear but might still be present only under certain conditions. Late in the season, when plants might have fewer resources available for reproduction and at the high and intermediate site where cold and dry environmental conditions are less favorable for the plants, the discrimination among distances was stronger.     

SEX RATIO,SEED PRODUCTION,BIOMASS ALLOCATION,AND THE COST OF MALE FUNCTION IN CUCURBITA FOETIDISSIMA HBK (CUCURBITACEAE)

In the gynodioecious plant Cucurbita foetidissima (Cucurbitaceae), females were common in all eight populations examined and made up 32% of adult plants. Females produced 1.5 (SE = 0.2) times as many seeds as did hermaphrodites. The observed difference in seed production alone is not great enough to explain the maintenance of females, especially at their current frequency. Females and hermaphrodites did not differ in number of nodes per stem, stems per plant, internode length, or size of leaves. Females produced more female biomass (fresh or dry weight) than hermaphrodites, but total investment in sexual biomass did not differ. Thus, the biomass of male flowers produced by hermaphrodites was about equal to the extra female biomass produced by females. The results support the existence of a trade-off between male and female reproduction.     

PATTERNS IN LEAF MORPHOLOGY AND PHOTOSYNTHESIS IN SHOOTS OF SASSAFRAS ALBIDUM (LAURACEAE)

Trees of Sassafras albidum (Nuttall) Nees. display leaves that fall into discrete categories of form. Unlobed leaves, with undissected margins, predominate at proximal and distal nodes of shoots, while lobed leaves are most common at intermediate nodes. In this study we investigated the hypothesis that shoots of sassafras respond to environmental changes over the growing season by generating predictable nodal patterns for leaf morphology and physiology. We recorded leaf shape category and measured leaf surface area, node number, chlorophyll content, nitrogen content, and photosynthesis in four trees. Over 1,000 measurements of photosynthesis were made in situ, using the LI-Cor LI 6200 portable photosynthesis system. The two trees growing under heavy shade had few lobed leaves (2.0% and 18.8%) and often had negative carbon balance, with positive net photosynthesis occurring during sun flecks. The two trees growing in more open conditions had many more lobed leaves (56.3% and 61.0%) and higher, less variable net photosynthetic rates. As indicated by chlorophyll and nitrogen contents, the highest leaf photosynthetic rates occurred at intermediate nodes (nodes six to nine), and this was shifted distally along the shoot during the growing season. Leaves at intermediate nodes also had the largest surface areas and the greatest frequency of lobing. In comparative experiments with models, lobing was shown to enhance free-convectional heat loss relative to unlobed leaves of the same surface area. Due to their large surface area, these leaves also have the highest rates of whole leaf photosynthesis. Under conditions of equivalent photosynthetically active radiation, lobed leaves had higher rates of net photosynthesis than did unlobed leaves. We conclude that shoots of sassafras produce groups of leaves with predictably different morphological and physiological specializations.     

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.     

EFFECTS OF SOIL NITROGEN ON POLLEN PRODUCTION,POLLEN GRAIN SIZE,AND POLLEN PERFORMANCE IN CUCURBITA PEPO (CUCURBITACEAE)

To determine the effects of soil nitrogen on pollen production, pollen size, and pollen performance, two cultivars of zucchini (Cucurbita pepo) were grown under two nitrogen regimes in an experimental garden. The two cultivars were true breeding for alternative alleles for a one gene trait, ovary color. The soil nitrogen treatment had a significant effect on most measures of reproductive output through the female function. The nitrogen treatment did not affect the number of staminate flowers per plant but did have an effect on the number of pollen grains per staminate flower and the mean pollen grain size. A pollen mixture experiment revealed that pollen produced by plants in the high nitrogen treatment sired significantly more seeds than pollen from low nitrogen plants. Moreover, we found that the high nitrogen pollen sired even a greater percentage of seeds in the region of the fruit (ovary) previously shown to be fertilized by the fastest growing pollen tubes. Thus, the difference in the number of seeds sired by pollen from the two nitrogen treatments is due to differences in pollen performance. We conclude that spatial heterogeneity in soil nitrogen can influence the paternity of seeds in a plant population.     

樟科木姜子属群的起源与演化

论述了樟科木姜子属群的起源与演化,认为该属群起源于古北大陆南部和古南大陆北部,以及古地中海周围热带地区,起源时间不早于中白垩纪。这一自然属群明显存在着平行进化情况,其核心的木姜子属和山胡椒属极可能是同地起源于共同祖先,我国南部至印度马来西亚可能是这两属的起源和分化中心,并从这里向热带美洲及大洋洲扩散。其余的属可能从这一核心演化出来。     

POTENTIAL MECHANISMS FOR SEX RATIO ADJUSTMENT IN MAMMALS AND BIRDS

Sex ratio skews in relation to a variety of environmental or parental conditions have frequently been reported among mammals and, though less commonly, among birds. However, the adaptive significance of such sex ratio variation remains unclear. This has, in part, been attributed to the absence of a low-cost physiological mechanism for sex ratio manipulation by the parent. It is shown here that several recent findings in reproductive biology are suggestive of many potential pathways by which gonadotropins and steroid hormones could interfere with the sex ratio at birth. And these hormone levels are well-known to be influenced by many parameters which have been invoked in correlating with offspring sex ratios. Hence, it is argued that the significant, but inconsistent sex ratio biases reported in mammalian and avian populations are coherent with current knowledge on reproductive physiology in those species. However, whether such variations can be viewed at as a consequence of physiological constraint or as adaptive sex ratio adjustment, has still to be determined.