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.     

INTRAPOPULATIONAL VARIATION IN POLLEN-MEDIATED GENE FLOW IN PLANTAGO LANCEOLATA L.

The extent of genetic variation in pollen-mediated gene flow distance was measured for a population of the wind-pollinated herb, Plantago lanceolata. Twenty-nine genotypes were collected from a natural population, clonally replicated, and grown to reproductive maturity in a greenhouse. Relative gene flow distances were measured for each replicate and genotype in a wind tunnel. Approximately five percent of the total variation in gene flow distance was attributable to variation among genotypes. Most of the remaining variation was attributable to differences between the early part of the growing season, when most of the flowering occurs, and the remainder of the season, when flowering is sparse. The rankings of the genotypes' gene flow distances showed significant concordance between the early-season measures and measures from later in the summer. There was no correlation between the average inflorescence height for a plant and the average gene flow distance for that plant. For analysis of pollen characteristics suspected as causal factors of variation in gene flow distance, the variation in gene flow distance was divided into two components: that due to environmental (seasonal) differences, and that due to the differences among genotypes within runs. Variation in both the buoyant properties of the pollen grains and their adhesiveness was significantly partially correlated with environmental variation in gene flow distance, while only the buoyant properties of the pollen grains were significantly partially correlated with among-genotype variation in gene flow distances.     

DYNAMICS OF GENE FLOW IN AN EXPERIMENTAL POPULATION OF CUCUMIS MELO (CUCURBITACEAE)

Gene flow in an experimental garden of Cucumis melo (Cucurbitaceae), the cultivated muskmelon, was measured by placing plants carrying a dominant gene (green cotyledons) in the center of an 18 × 18-m field of recessive plants (yellow cotyledons). At the end of the growing season, all fruits were collected, and seeds planted for analysis of the progeny. The 123 fruits yielded 41,875 seedlings whose genotypes were recorded. Gene flow decreased with increasing distance from the central plot, while the variance increased. The decrease in frequency of the green gene did not follow a leptokurtic pattern over the distance measured, and there was a marked asymmetric and patchy pattern of gene flow in the field. The directionality contributed to a bimodal pattern of gene frequencies among fruits at the edges of the field, some fruits with no green seedlings, some with many. Bumblebee movements were recorded during 4,296 flower visits in an identically arrayed melon field. Bees usually moved to very nearby plants and visited only a few flowers on any one plant. This suggests that pollen carryover from the central patch out was quite high in the test garden. These findings show that local gene flow patterns in plant populations may be complex, and do not always follow regular and symmetrical models. The complex arrangement of progeny genotypes, when acted upon by local selective forces, may contribute to small-scale differences often seen in plant populations.     

HETEROCHRONY AND HETEROBLASTIC LEAF DEVELOPMENT IN TWO SUBSPECIES OF CUCURBITA ARGYROSPERMA (CUCURBITACEAE)

To date, reports of paedomorphosis at the whole plant or shoot level have been loosely based on whole plant form or on the sequence of leaf shapes produced along the shoot (heteroblasty). However, interpreting the significance of heterochrony in the evolutionary loss or gain of heteroblasty based on mature leaf forms assumes that all leaves with the same shape arose through very similar modes of organogenesis. This study examines this assumption in two subspecies of Cucurbita argyrosperma, one that is wild and heteroblastic and a second that is cultivated and not markedly heteroblastic. All leaves of the cultivar are visually similar to early leaves of the wild subspecies. The cultivar is considered to be the progenitor of the wild subspecies. Scanning electron microscopy and allometry of developing leaves showed that at early nodal positions along the primary shoot, leaf development in both subspecies was similar. At later nodal positions, very young leaves of both subspecies were more similar to each other than to leaves at earlier nodal positions within the same plant at the same stage of development. This early similarity was masked in the mature shapes of later leaves due to subsequent differences in allometric growth. Thus a simple hypothesis of paedomorphosis in which the early leaf form in the progenitor is simply reiterated at later nodal positions in the cultivar is not supported by patterns of leaf development.     

INTERSPECIFIC GENE FLOW IN CUCURBITA: C. TEXANA VS. C. PEPO

The potential for interspecific genetic exchange was examined by monitoring flowering patterns, pollinator movement, and gene flow among experimental populations of the Texas gourd (Cucurbita texana) and cultivars of Cucurbita pepo. While flowering patterns and pollinator movement tended to maximize self-pollination and local gene exchange, movement of effective pollen exceeded 1,300 m. This movement, mediated by the solitary bee Xenoglossa strenua and monitored by tracking allozyme variants, produced interspecific hybrids in 5% of the progeny from experimental plants. Interspecific gene exchange occurred in either direction with either species serving as staminate or pistillate parent. No obvious constraints to gene flow among plants representing C. texana and distinctive cultivars (vars. ovifera, medullosa, melopepo) of C. pepo were detected. Genetic exchange among different species and cultivars is enhanced by the foraging behavior of Xenoglossa. Multiple visits to either staminate (pollen carryover) or pistillate (multiple pollinations) flowers often result in the deposition of mixed pollen on receptive stigmas. The wild type (C. texana) can donate and receive effective pollen when growing under both weedy and natural conditions. The observed lack of interspecific reproductive isolation supports treatment of cultivars and wild types as a single species and, in conjunction with available data concerning temporal/geographical relationships among bees, squash, gourds, and humans in eastern North America, suggests the possibility of long-term genetic interaction between wild types and domesticates.     

GENE FLOW IN LICHENS

Experimental studies of the evolutionary biology of lichen fungi have been hampered by massive difficulties of in vitro culture and artificial crosses are still not possible. Gene flow in these organisms is demonstrated here for the first time by the analysis of secondary products in the progeny of individuals from natural populations of mixed chemotypes of the Cladonia chlorophaea complex. All of the chemotypes in this study have been interpreted as distinct sibling species. In the Appalachian Mountains of North Carolina, however, the grayi and merochlorophaea chemotypes are found to belong to a single interbreeding populations that is reproductively isolated from the cryptochlorophaea chemotype. In the Coastal Plain, the cryptochlorophaea chemotype hybridizes with the local endemic perlomera chemotype. This study has major impact for species concept in lichens because consideration of neither morphologial tendencies nor biogenetic relationships of the secondary products could have predicted its result.     

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.     

GENETIC STRUCTURE AND MALE-MEDIATED GENE FLOW IN THE GHOST BAT (MACRODERMA GIGAS)

The Australian ghost bat is a large, opportunistic carnivorous species that has undergone a marked range contraction toward more mesic, tropical sites over the past century. Comparison of mitochondrial DNA (mtDNA) control region sequences and six nuclear microsatellite loci in 217 ghost bats from nine populations across subtropical and tropical Australia revealed strong population subdivision (mtDNA φ_ST = 0.80; microsatellites UR_ST = 0.337). Low-latitude (tropical) populations had higher heterozygosity and less marked phylogeographic structure and lower subdivision among sites within regions (within Northern Territory [NT] and within North Queensland [NQ]) than did populations at higher latitudes (subtropical sites; central Queensland [CQ]), although sampling of geographically proximal breeding sites is unavoidably restricted for the latter. Gene flow among populations within each of the northern regions appears to be male biased in that the difference in population subdivision for mtDNA and microsatellites (NT φ_ST = 0.39, UR_ST = 0.02; NQ φ_ST = 0.60, UR_ST = ?0.03) is greater than expected from differences in the effective population size of haploid versus diploid loci. The high level of population subdivision across the range of the ghost bat contrasts with evidence for high gene flow in other chiropteran species and may be due to narrow physiological tolerances and consequent limited availability of roosts for ghost bats, particularly across the subtropical and relatively arid regions. This observation is consistent with the hypothesis that the contraction of the species' range is associated with late Holocene climate change. The extreme isolation among higher-latitude populations may predispose them to additional local extinctions if the processes responsible for the range contraction continue to operate.     

GENETIC VARIATION AND ESTIMATES OF GENE FLOW IN CLARKIA SPECIOSA SUBSP. POLYANTHA (ONAGRACEAE)

Estimates of interpopulational gene flow and the levels and distribution of genetic variation in Clarkia speciosa subsp. polyantha were obtained using enzyme electrophoresis. Eight enzymes encoded by 17 loci were analyzed. Nei's mean genetic identity was 0.96, indicating little genetic divergence among populations. Gene diversity statistics also suggest little heterogeneity among populations. Interpopulational gene flow, estimated according to Slatkin (1985), was fairly high, Nm = 3.9, probably accounting for the lack of differentiation among populations.     

ADAPTIVE POPULATION DIVERGENCE IN CRYPTIC COLOR-PATTERN FOLLOWING A REDUCTION IN GENE FLOW

Adaptive population divergence is often driven by divergent natural selection, but can be constrained by the homogenizing effect of gene flow between populations. Indeed, a common pattern in nature is an inverse correlation between the degree of adaptive phenotypic divergence between populations and levels of gene flow between populations. However, there is essentially no experimental data on whether this correlation arises because gene flow constrains adaptation or, conversely, because adaptive divergence causes barriers to gene flow (ecological speciation). Here, I report increased adaptive divergence in cryptic color pattern between a pair of Timema insect populations following an experimental reduction in between-population gene flow. The reduction in gene flow arose due to a natural experiment, and thus was not replicated at a second site. However, temporal replication of the trends among six generations of data, coupled with a lack of increased adaptive divergence for two other population pairs where gene flow was not manipulated (i.e., control sites), argues that the results did not arise by chance. Estimates of dispersal ability and population size further support reduced gene flow, rather than increased genetic drift, as the cause of divergence. Thus, the findings provide experimental evidence that gene flow constrains adaptation in nature.     

SEEDLING MORPHOLOGY IN MARAH (CUCURBITACEAE) RELATED TO THE CALIFORNIAN MEDITERRANEAN CLIMATE

Studies in reproductive ecology were made in indigenous, western American plants in the genus Marah (Cucurbitaceae), with particular attention given plants of M. oreganus occurring in the Berkeley Hills near San Francisco Bay in California. These tuberous perennials produce capsular fruits on their annual aboveground shoots; the fruits dehisce in early summer, each one exposing about three large seeds with an average seed weight of 1.05 g. The embryo of a M. oreganus seed has two thick and fleshy cotyledons packed with protein granules. The embryonic axis, with shoot and root apices, is ca. 0.5-1.0 mm long, roughly ½0 or less the length of the seed. In the Berkeley Hills dispersal of the seeds is accomplished by nocturnal rodents, after which germination begins with the fall rains and cooler temperatures of November and December. Instead of a radicle emerging first from the seed at germination, the minute radicle and epicotyl are pushed or carried far out of the seed, down into the soil, by the elongating bases of the cotyledons. These cotyledon bases, or petioles, are fused, and as they elongate they form a hollow tube that bears the embryonic axis at its extreme tip. The cotyledonary petiole tube ceases elongation by January, when it may be 5-25 or more cm long in a seedling of M. oreganus. Then, from its tip, the radicle grows downward and the epicotyl upward—up the hollow petiole tube. The green shoot (epicotyl) reaches the soil surface by early March in this area, completes the first season's growth, and dries up by late May, when the arid summer season is beginning. But even before the epicotyl grows out of the petiole tube and above ground, the seedling's hypocotyl begins to enlarge, forming a tuber. The fleshy cotyledon blades remain in the seed coat below ground, and some food from the blades is transferred to the tuber that produces shoots in the following growing seasons. This pattern of germination and seedling establishment is now known for species of Marah and for a very few other dicotyledonous plants, all of them growing mainly in areas of hot and dry habitat that are generally referred to as having Mediterranean climate. This elongation of the fused hypogeal cotyledons is considered a complex adaptation in dicotyledons that helps ensure fast and successful seedling establishment in seasonally arid areas such as “Mediterranean” California.     

MONOMORPHISM,REDUCED GENE FLOW,AND CLEISTOGAMY IN RARE AND COMMON SPECIES OF LESPEDEZA (FABACEAE)

Population genetic structure was analyzed in the rare, native prairie legume Lespedeza leptostachya Engelm. and in the widespread L. capitata Michx. Both species produce a mixture of chasmogamous and cleistogamous flowers. Allozymes were analyzed for 32 loci from 224 individuals from 12 populations of L. leptostachya and for 34 loci in 291 individuals from 12 populations of L. capitata. L. leptostachya is entirely monomorphic at all loci studied, while L. capitata shows strong among-population differentiation for the limited variation that occurs in that species. Allozyme data suggest that the level of gene flow among populations of L. capitata is very low, and that very low levels of outcrossing are effected by the chasmogamous flowers in L. capitata.     

赤■属的系统学研究

采用比较形态学方法,本文详细地描述了赤属植物根、根状茎、茎、卷须、花序、花萼、花冠、雄蕊群、雌蕊群、果实、种子以及毛被的形态特征,并讨论了某些形态学性状的系统学意义。本文首次描述和记录了该属植物的传粉生物学特征以及传粉媒介,并讨论了造成该属某些种结实率低的原因。本文还根据现存赤属及其近缘属植物的分布式样以及它们的属下分类群的系统演化式样,确定了中国的横断山系和大巴山系地区是该属植物的现代分布中心和分化中心,而中国云南南部和中南半岛北部地区的季节性山地雨林是该属植物的起源中心。在综合大量资料的基础上,本文对赤属作了分类学修订。确认该属有２２种,２变种,归并了２种和１０变种。