新疆郁金香营养生长、个体大小和开花次序对繁殖分配的影响

植物有性繁殖与资源分配的关系研究对于揭示植物生活史特征及繁育系统进化具有重要意义。新疆郁金香(Tulipa sinkiangensis)是新疆天山北坡荒漠带特有的一种多年生早春短命植物。在自然生境中,该物种仅以有性繁殖产生后代,每株能产生1-8朵花,且不同植株上的花数及果实数以及花序不同位置上的花与果实大小明显不同。本文通过对新疆郁金香有性繁殖与营养生长及植株大小的关系以及花序中不同位置花及果实间的资源分配研究,旨在揭示营养生长、个体大小及开花次序对其繁殖分配的影响。结果表明:在开花和果实成熟阶段,新疆郁金香植株分配给营养器官(鳞茎和地上营养器官)与繁殖器官的资源间均存在极显著的负相关关系(P<0.01),说明其植株的营养生长与生殖生长间存在权衡关系。多花是新疆郁金香的一个稳定性状,其植株上花数目、花生物量、果实生物量和种子数量与植株生物量间均呈极显著的正相关关系(P<0.01),说明新疆郁金香植株的繁殖分配存在大小依赖性。在具2-5朵花的新疆郁金香植株中,花序内各花的生物量、花粉数和胚珠数、结实率、果实生物量、结籽数、结籽率及种子百粒重按其开花顺序依次递减,说明花序内各花和果实的资源分配符合资源竞争假说。植株通过减少晚发育的花或果实获得的资源来保障早发育的花或果实获得较多的资源,从而达到繁殖成功。     

Fruit set,fruit reduction,and fruiting strategy in Cornus sanguinea (Cornaceae)

We studied various aspects of the fruiting biology of Cornus sanguinea (Cornaceae), a fleshy-fruit-producing deciduous shrub, in four populations in northwest Spain. One population was studied over a 5-yr period (1989-1994), and the remaining populations in 1994 only. Fruit-set level varied among years (range 11-18%) and among populations (range 8-22%), but was in all cases low. Within plants, fruit-set level did not vary significantly among inflorescences, indicating that inflorescence fruit set is independent of inflorescence position and inflorescence, phenology. To investigate the function of surplus flowers, we carried out flower removal experiments. Inflorescence fruit-set level was unaffected by removal of up to =75% of flowers. These results suggest that Cornus sanguinea regulates its fruit-set level via plasticity in the number of fruits aborted: if flower mortality has been high, fewer fruits will be aborted. Within the inflorescence, surplus flowers thus act as insurance against flower loss.     

A REEXAMINATION OF THE POLLEN-DONATION HYPOTHESIS IN AN EXPERIMENTAL POPULATION OF ASCLEPIAS EXALTATA

The evolution of large floral displays in hermaphroditic flowering plants has been attributed to natural selection acting to enhance male, rather than female, reproductive success. Proponents of the “pollen-donation hypothesis” have assumed that maternal resources, rather than levels of effective pollination, limit fruit set. We investigated the pollen-donation hypothesis in an experimental population of poke milkweed, Asclepias exaltata, where effective pollination did not limit fruit set. Specifically, we examined the effects of flower number per plant, and flower number per umbel on male reproductive success (number of fruits sired) and female reproductive success (number of fruits matured). In 1990, a paternity analysis was performed on fruits collected from 53 plants whose inflorescences were not manipulated. Flower number per plant was significantly correlated with male success, but not with plant gender. Flower number per plant was also significantly correlated with female success, but umbel number and stem number per plant together explained more than half (58%) the variation in female success. The percentage of fruit set was not significantly correlated with flower number per plant. Plants with large floral displays did not disproportionately increase in male reproductive success, relative to female success, as predicted by the pollen-donation hypothesis. In 1991, the effect of flower number per umbel on male and female reproductive success was investigated. Flower number per umbel was manipulated on four umbels per plant by removing flowers to leave 6, 12, or 18 flowers in each umbel. Plants with the largest umbels effectively pollinated twice as many flowers on other plants, but produced only 1.35 times as many fruits as plants with 6 and 12 flowers per umbel. Relative maleness of plants with large umbels was nearly twice that of small and medium umbels. Although these observations are consistent with the pollen-donation hypothesis at the level of umbels, they are problematic, because much of the variation in flower number per umbel exists within, rather than among, plants in natural populations. Thus, plants consist of both reproductively male (large) and female (small) inflorescences, which act to increase total reproductive success. It is therefore inappropriate to explain the evolution of large floral displays in milkweeds solely in terms of potential male reproductive success.     

REPRODUCTIVE OUTPUT AND BIOMASS ALLOCATION IN SESBANIA EMERUS IN A TROPICAL SWAMP

This study examined the correlation of moisture, reproductive phenology, density of mature plants, and herbivory of apical meristems with the morphology and reproductive output of Sesbania emerus, an annual legume growing along a moisture gradient in a swamp in Guanacaste Province, Costa Rica. It also determined how biomass allocation varied and how it was related to reproductive output of plants growing along the moisture gradient within the swamp. Morphological changes included production of more stems and branches in response to herbivory, more and higher prop roots and more aerenchyma as water depth increased, and greater stem diameter in lower densities. Plant height varied greatly within a site, but not among habitats. Plants began to produce flowers and fruits later in wet sites than in dry sites. Reproductive output was generally more sensitive to environmental variables than was plant size. Fruit number and plant height were positively correlated for almost every treatment. Greater fruit and seed production were correlated with drier sites, earlier phenology, and lower density, but not with herbivory. Total biomass accumulation did not vary among moisture sites, but root production appeared to occur at the expense of reproductive output in the wetter sites. Plants in the wetter sites had both a greater percent and a greater absolute amount of biomass in roots, and a lower percent and a lower absolute amount of biomass in fruits and seeds than plants in drier sites. The root: shoot ratio was nearly five times higher in the wet than the dry site. Seed number per plant ranged from a mean of 6,800 at the wet site to a mean of 16,878 at the dry site. If this striking phenotypic variation in reproductive output and biomass accumulation has a genetic basis, the possibility of ecotypic differentiation exists in S. emerus.     

Size-dependent pollination efficiency in Anchusa officinalis (Boraginaceae): causes and consequences

Summary Bumblebees foraging on the self-incompatible Anchusa officinalis fly between near neighbour plants and between near neighbour inflorescences within plants. Although many-flowered plants attracted most bumblebees these plants received fewer visits on a per flower basis than smaller plants, and each bumblebee visited a smaller proportion of the flowers. The calculated effective visitation rate per flower was highest on plants of an intermediate size. If pollen-carryover was assumed to be limited the most efficient plant was predicted to be smaller since the proportion of fertilized flowers per bumblebee visit is expected to decrease further on the largest plants in relation to the total flower number. These predictions were tested by measuring fruit-set in the field. The percentage fruit-set decreased with plant size at all sizes that were investigated. That the most efficient plant was small indicates that pollen-carryover was indeed limited. However, the low percentage fruit-set associated with large size did not present a serious problem since the total estimated seed production per plant still increased with size. Selection favoring smaller plants may be low or absent in Anchusa.     

Effects of population size on reproductive success of the endangered and endemic species Primula merrilliana

The reproductive ability related to the population size of the endangered and endemic species Primula merrilliana Schltr.was investigated.In 26 populations observed,only four contain more than 500 flowering individuals,whereas most of them(53.8%) consist of less than 100 flowering individuals.Though the ratio of pin and thrum plants keeps its balance at 1:1 for all populations,the frequency of pin and thrum flowers was significantly biased in most small populations.Population size strongly affected reproductive success; plants in small populations produced significantly fewer flowers and fruits per plant and fewer seeds per fruit,and therefore fewer seeds per plant.The floral density was another important factor that influenced the reproductive success of P.merrilliana,because four main reproductive success parameters (fruits per plant,seeds per fruit,seeds per plant,and the proportion of flowers setting fruit) were all positively correlated with floral density.The size of plants and the number of leaves per plant (measure of habitat quality) were not influenced by the variation of population size,suggesting that the reduced fecundity in small populations may not be a consequence of lower habitat quality,inbreeding depression and pollen limitation as a result of less attractiveness in small populations are therefore likely explanations for these patterns.     

THE RELATIVE COSTS OF SELF- AND CROSS-FERTILIZED SEEDS IN IMPATIENS CAPENSIS (BALSAMINACEAE)

By using a generally applicable technique that involves monitoring the development and survivorship of flowers and seed capsules, I estimated the material and energetic costs of producing self- and cross-fertilized seeds in Impatiens capensis. All flowers and fruits on six plants were censused intensively for the two-month period of reproduction. Cleistogamous (selfing) flowers ripened seed in about 24 days, compared to about 36 days for the chasmogamous (outcrossing) flowers. In terms of dry weight, selfed seeds cost about two-thirds as much as outcrossed seeds: 12.4 versus 18.4 mg dry weight per seed. When adjusted to the currency of calories, and including an independent estimate of pollen and nectar production in outcrossing flowers, I estimate the costs to be about 65 and 135 calories per selfed or outcrossed seed. Sources of error include the accuracy of the estimates of flower and fruit weight, and possible differences among the developmental stages in respiratory costs. The cost discrepancy implies that outcrossed seeds should possess a countervailing fitness advantage large enough to offset their greater energetic cost.     

TEMPORAL VARIATION OF FLOWER AND FRUIT SIZE IN RELATION TO SEED YIELD IN CELANDINE POPPY (CHELIDONIUM MAJUS; PAPAVERACEAE)

Ecological and evolutionary studies typically consider variation in single reproductive characters in isolation, without considering how they might be correlated with other reproductive and vegetative characters. In our study, we examined temporal patterns of variation and correlation in flower diameter and fruit length during a reproductive phase in two Massachusetts populations of the herb, Chelidonium majus. We also examined the relationships of such variation to measurements of seed yield components (mean seed weight and number per fruit) and aspects of plant vegetative size. Most of the variation in the sizes of reproductive characters occurred within individual plants, instead of among plants or between populations. Flower and fruit sizes as well as seed number per fruit declined significantly during the season in both populations. Only mean seed size per fruit was relatively stable for individual plants in both populations. Conserving resources by a gradual reduction in the size of reproductive characters over the season may be a strategy for maternal plants to continue seed production. The strong, persistent patterns of correlation between certain characters, such as flower and fruit size, in spite of extensive phenotypic plasticity, was interpreted as indirect evidence for developmental correlation. Furthermore, vegetatively larger plants produced not only more flowers and fruits, but also consistently larger flowers and fruits. The results emphasize that variation in fitness characters, such as seed size and number, should not be viewed in isolation from vegetative characters, flower, and fruit sizes in ecological and evolutionary studies, if the goal is to understand the mechanisms of natural selection in wild populations.     

Changes in growth and photosynthetic characteristics of Ocimum sanctum under growth regulator treatments

A field experiment was conducted to investigate the effect of growth regulators on growth characteristics such as root length, shoot length, total leaf area, number of inflorescence per plant, number of flower per inflorescence, whole plant fresh weight and whole plant dry weight. Photosynthetic characteristics were also analyzed based on the same experiment. For this, various photosynthetic pigment contents such as chlorophyll, carotenoid, anthocyanin and xanthophyll content were calculated. The conventional growth regulator abscisic acid (ABA) and non-conventional growth regulator triazole compound paclobutrazol (PBZ) were used. Root length increased due to growth regulator treatment, but shoot length decreased. Leaf area was decreased due to growth regulator treatment. The number of inflorescence increased in ABA treated plants, but it was decreased in PBZ treated plants. In ABA treated plants, the number of flowers per inflorescence was increased. In PBZ treated plants the number of inflorescence was reduced. The whole plant fresh weight (FW) and dry weight (DW) were increased in ABA and PBZ treated plants. There was an increase in chlorophyll content in growth regulator treated plants compared to control, and it was more in PBZ treated plants. The carotenoid content was also increased in ABA and PBZ treated plants.     

Do local conspecific density and floral display size influence fruit set via pollinator visitation in Orchis militaris?

Plant density varies naturally, from isolated plants to clumped individuals, and this can influence pollinator foraging behaviour and plant reproductive success. In addition, the effect of conspecific density on reproduction may depend on the pollination system, and deceptive species differ from rewarding ones in this regard, a high density being often associated with low fruit set in deceptive plants. In our study, we aimed to determine how local conspecific density and floral display size (i.e. number of flowers per plant) affect fruit set in a deceptive orchid (Orchis militaris) through changes in pollinator visitation. We measured fruit set in a natural population and recorded pollinator abundance and foraging behaviour within plots of different O. militaris densities. Detailed data were recorded for the most abundant potential pollinators of O. militaris, i.e. solitary bees. Floral display size was negatively correlated to fruit set in medium‐density plots, but uncorrelated in low‐ and high‐density plots. Plot density had no effect on solitary bee abundance and visitation, which may be due to low pollinator abundance within the study site. The proportion of visited flowers per inflorescence was negatively influenced by floral display size, which is in line with previous studies. In addition, solitary bees spent decreasing time in successive flowers within an inflorescence, and the time spent per flower was negatively affected by ambient temperature. Our results suggest that pollinator behaviour during visitation is poorly linked to pollen deposition and reproductive success in O. militaris.     

The role of termites in the decomposition of wood and leaf litter in the Southern Guinea savanna of Nigeria

Summary The number of flowers produced by inflorescences of Yucca whipplei (Agavaceae) consistently exceeds the number of fruits produced by about one order of magnitude. To determine the factors responsible for low fruit set, the relation between pollinator availability, the amount of resources spent on reproduction (as indicated by inflorescence size), and the number of fruits matured was studied during 1978 and 1979 at 18 locations in chaparral, coastal sage scrub, and desert scrub communities of southern California.The following results support the conclusion that pollinators do not usually limit fruit production in Yucca whipplei. Rather, fruit production is limited by the amount of resources available to support developing fruits. (1) Fruit production is positively correlated with inflorescence size both within and between populations. The average size of inflorescence for a population is an excellent predictor of mean fruit production. Furthermore, 54% of the total variance in fruit production of individual plants can be explained by inflorescence size. (2) In contrast, although fruit production within most populations is positively correlated with an index of the number of pollinator visits to an inflorescence, the relative abundance of pollinators for a population is a poor predictor of mean fruit production, and only 9% of the total variance in fruit production can be explained by the visitation index. Furthermore, at four sites studied for two years, there was little change in average inflorescence size or fruit production from 1978 to 1979, despite large differences in relative abundance of pollinators at each of the sites. (3) Based on geographic proximity, and physiographic and vegetational similarities, study sites were grouped into regional clusters. Both inflorescence size and fruit production varied considerably between regions. Of the total variation in fruit production, 27% can be attributed to differences between regions. Most of this variation is the result of regional differences in inflorescence size, which in turn influence fruit production.Why does Yucca whipplei produce such large inflorescences if so few fruits can be supported? Two relevant hypotheses are discussed: (1) the floral display is the result of selection for pollen dissemination at the expense of fruit set; and (2) the floral display is the result of selection for a bet-hedging strategy either to increase the probability of adequate pollination when pollinators are unusually rare, or to allow individuals to support more fruits when resources are unusually abundant.     

荒漠植物准噶尔无叶豆结实、结籽格局及其生态适应意义

连续2a采用野外记录的方式对荒漠植物准噶尔无叶豆的结实、结籽格局进行了比较研究。结果表明:植株内花序生成格局表现为不同大小的花序在总花序中的比重与其结实率存在显著正相关,其中含有中等花数目的花序所占的比重及其结实率均最高,此种分布格局能够最大程度的保证繁殖成功;果序内果实生成格局表现为单花着生位置(从近柄端算起)与其结实量占总结实量的比例成线性负相关,花序基部的结实比例最高,顶部结实比例近为0;荚果内种子生成格局表现为:中间位置的胚珠败育率最低。另外,年际间荚果内仅有(1.08±0.03)粒和(1.07±0.03)粒种子能够完全成熟,此种结籽格局是准噶尔无叶豆保证后代质量的最佳策略。     

Conservation of microRNA-mediated regulatory networks in response to copper stress in grapevine

Jatropha curcas L. as a bio energy plant belonging to the Euphorbiaceae family is gaining progressive importance over the last years. In 2012 and 2014 field experiments were carried out to assess the effects of cytokinins 6-Benzyladenine (BA) and Forchlorfenuron (CPPU) acting as plant growth regulators (PGRs). Number of flowers per inflorescence, female-to-male ratio of flowers, fruits per infructescence, fruiting rate, number of seeds per fruit, seed size and weight as well as seed oil content were determined. It was suggested that concerning effectiveness of exogenous application of PGRs the developmental stage of flower is very important. Due to that, special interest was laid on the age of inflorescences at the time of application. Our experiments revealed a strong dependence of cytokinin effectiveness on the developmental stage of flowering. So treatment of plants with 6-Benzyladenine at the beginning of flower formation (stage 1) significantly increased the number of male and female flowers per inflorescence, whereas treatment at an advanced flowering stage (stage 2) or at the stage of fully developed flowers (stage 3) had only slight or no effects. In contrast, fruit retention was progressively increased by treatment in stage 2 and 3. Application of Forchlorfenuron significantly increased female-to-male ratio in stage 1 flowers but showed no effects on stage 2 and 3. 6-Benzyladenine as well as Forchlorfenuron showed equal effects on number of fruit inflorescences treated in stage 1. Our results show a significant decrease in seed weight due to BA- and CPPU-application while kernel weight remained stable. Concerning fruits, clusters and oil yield per tree, BA-application showed significant increasing effects. This study indicates that 6-Benzyladenine and Forchlorfenuron can improve yield affecting parameters in J. curcas when the phenological stage of flowering at time of application is precisely considered.     

Changes in growth and photosynthetic characteristics of Ocimum sanctum under growth regulator treatments

A field experiment was conducted to investigate the effect of growth regulators on growth characteristics such as root length,shoot length,total leaf area,number of inflorescence per plant,number of flower per inflorescence,whole plant fresh weight and whole plant dry weight.Photosynthetic characteristics were also analyzed based on the same experiment.For this,various photosynthetic pigment contents such as chlorophyll,carotenoid,anthocyanin and xanthophyll content were calculated.The conventional growth regulator abscisic acid (ABA) and non-conventional growth regulator triazole compound paclobutrazol (PBZ) were used.Root length increased due to growth regulator treatment,but shoot length decreased.Leaf area was decreased due to growth regulator treatment.The number of inflorescence increased in ABA treated plants,but it was decreased in PBZ treated plants.In ABA treated plants,the number of flowers per inflorescence was increased.In PBZ treated plants the number of inflorescence was reduced.The whole plant fresh weight (FW) and dry weight (DW) were increased in ABA and PBZ treated plants.There was an increase in chlorophyll content in growth regulator treated plants compared to control,and it was more in PBZ treated plants.The carotenoid content was also increased in ABA and PBZ treated plants.     

SIZE-DEPENDENT GENDER CHANGE IN GREEN DRAGON (ARISAEMA DRACONTIUM; ARACEAE)

Green dragon (Arisaema dracontium; Araceae) is a perennial woodland herb capable of switching gender from year to year. Small flowering plants produce only male flowers but when larger they produce male and female flowers simultaneously. Distinct male and monoecious phenotypes (referred to hereafter as plants) share a single underlying cosexual genotype. Four populations in southern Louisiana were sampled to determine frequencies and size distributions of male and monoecious plants, and to determine the relationship of plant size with male and female flower production in monoecious plants. Male plants were significantly smaller than monoecious plants and made up 34%–78% of flowering plants within populations. Flower number (average = 120) was weakly positively correlated with size. Monoecious plants produced an average of 169 flowers (90 female) and had 100% fruit set, with individual berries containing an average of 2.5 ovules and 1.3 filled seeds. Male flower number was negatively correlated, and female flower number positively correlated, with basal stem diameter. Extrapolation of regression slopes suggested that green dragon should become completely female at a size 20% larger than the largest plant observed in this study. A simple model of inflorescence development is presented to illustrate how the reproductive system of green dragon is related to that of jack-in-the-pulpit (A. tnphyllum), which exhibits a more distinct switch between male and female phenotypes.     

Aridity mediates the effect of wood extraction on the reproductive output of an endemic disturbance-adapted woody species (Cenostigma microphyllum,Leguminosae) in the Caatinga dry forest

Chronic anthropogenic disturbances and climate change have been recognized as drivers of biological reorganization across human-modified tropical landscapes, also negatively affecting the reproductive output of some plant species. Here, we investigated to what extent these drivers affect the reproductive output of Cenostigma microphyllum, a disturbance-adapted tree species endemic to the Caatinga dry forest. The production of flowers/inflorescence, fruits and seeds was estimated for 105 plants (≥3 cm diameter at soil height) across 11 forest stands (20 × 50 m each), covering gradients of chronic disturbances (e.g. goat and livestock herbivory; wood extraction; and removal of non-timber forest products) and aridity at the Catimbau National Park. We documented that when the drivers were analysed in isolation, the fruit-set was positively associated with increased wood extraction, and the fruit-set and total number of seeds/plots were reduced under increasing aridity. We also verified a complex interaction between wood extraction and aridity emerged. In forest stands, the combination of high levels of aridity and wood extraction leads to a decrease in the fruit-set and total number of seeds/plots. Conversely, the fruit-set and total number of seeds/plots increased in stands exposed to lower aridity but high levels of wood extraction. Our results suggest that chronic disturbances and aridity affect plant fitness in a complex way including both negative and positive effects on attributes related to plant reproductive performance, which could result in both proliferation and population decline in the same landscape. Although C. microphyllum benefits from anthropogenic disturbances and is considered a disturbance-adapted species, it apparently does not benefit from increases in aridity, which is an expected future scenario for the Caatinga dry forest.     

Competition for assimilates and fruit position affect fruit set in indeterminate greenhouse tomato

Localization and characterization of fruit set in winter tomato crops was investigated to determine the main internal and external controlling factors and to establish a quantitative relationship between fruit set and competition for assimilates. Individual fruit growth and development was assessed on a beef tomato cultivar during the reproductive period (first nine inflorescences). A non-destructive photograph technique was used to measure fruit growth from very early stages of their development and then calliper measurements were made on big fruits. From these measurements we determined the precise developmental stage at which fruit growth stopped. Fruit potential growth, which is defined as the growth achieved in non-limiting conditions for assimilate supply, was also assessed by this method on plants thinned to one flower per inflorescence. The latter was used to calculate the ratio between actual and potential growth, which was found to be a good index of the competition for assimilates. Time lags of fruit set were observed mainly on distal organs. When more than three flowers were left on each inflorescence, distal organs developed at the same time as proximal organs of the following inflorescence. Consequently they were submitted to a double competition within one inflorescence and among inflorescences. It was shown that, what is commonly named 'fruit set failure', is not an irreversible death of the organ and that a small fruit could resume growth after a delay of several weeks as soon as the first fruits ripened and thus ceased to compete for assimilates. In that case proximal fruits resumed growth before distal ones. The delayed fruits contained only few seeds but a germination test confirmed that fertilization took place before fruit set failed. Competition for assimilates was calculated during plant development by the ratio between actual and potential fruit growth. Potential growth of proximal fruits was strongly dependent on the position of the inflorescence on the stem, whereas potential growth of distal fruits was lower than or equal to that of proximal fruits of the same inflorescence and it was independent on the inflorescence position. We took into account both inflorescence and fruit positions to establish a quantitative relationship between fruit set of individual inflorescences and the ratio between actual and potential fruit growth.     

Factors constraining fruit set in Mandevilla pentlandiana (Apocynaceae)

The reproductive success of Mandevilla pentlandiana was studied to disclose its reproductive strategy, and to determine the links between nectar production, breeding system, fruit set and inflorescence size. The plant produces many inflorescences with a large number of flowers but initiates few fruits (9%). This vine is self-compatible but not autogamous. Given that no significant differences could be detected considering many traits (ripe and abortive fruit sets, fruit quality, and seedling survival) between the pollination treatments (self-, cross-and natural-), the low natural fruit set was not related to pollen limitation. Fruits were not distributed at random within inflorescences (earlier fruits had the highest probability of maturation) but there were no significant differences in fruit quality according to different fruit positions. Conversely, the time of fruit initiation influenced most of the fruit-traits. Many developing fruits were aborted (20%). An increase in the probability of abortion was detected when the whole inflorescence was hand pollinated. In addition, a positive correlation was detected between the abortions and the number of ripe fruits which developed before them. Looking at our data from an evolutionary perspective, we argue that a theoretical inflorescence size, corresponding to the intersection point between the mean values of fruit number and fruit set per inflorescence, can be assumed to indicate the optimum inflorescence size that maximizes equally both female and male functions. Comparison between the theoretical and the observed mean inflorescence size suggests, that for M. pentlandiana , pollen donation may be the primary evolutionary factor behind excess flowers.