Trade-offs between flower number and investment to a flower in selfing and outcrossing varieties of Impatiens hypophylla (Balsaminaceae)

Floral resource allocation was compared on a whole-plant basis between two varieties of Impatiens hypophylla that differ in flower size. There were significant negative correlations between flower number and investments to a flower at both the within-population and between-variety levels. In individual flowers, var. hypophylla with larger flowers invested significantly more resources to male and pollinator-attractive functions, whereas investments to female function did not differ between the varieties. In experimental populations placed in the field, pollinators preferred the larger flowers of var. hypophylla even within the same habitat of var. microhypophylla, which has smaller flowers. There was a significant lack of observed heterozygosity only in var. microhypophylla. Thus, the outcrossing variety had more attractive but fewer flowers, while the selfing variety had less attractive but more abundant flowers.     

Chromosome evolution in Himalayan Impatiens (Balsaminaceae)

AKIYAMA, S., WAKABAYASHI, M. & OHBA, H., 1992. Chromosome evolution in Himalayan Impatiens (Balsaminaceae). Chromosome numbers and karyotypes have been investigated in species of Himalayan Impatiens . In addition to confirming previous chromosome counts, the presence of a tetraploid taxon ( I. exilis) is revealed. In central and east Nepal species with x = 9 are more common than those with other basic numbers and this number is shown to be one of the most frequent numbers in the genus. Most species with x = 9 have a bimodal karyotype. The species relationships are discussed.     

Comparisons of genetic diversity in a narrow endemic, Impatiens hypophylla, and in its widespread congener, I. textori (Balsaminaceae)

Genetic diversity in two species of Impatiens in Korea was investigated using starch gel electrophoresis. Impatiens hypophylla is a rare species endemic to southwestern Japan and southeastern Korea, whereas I. rextori is widespread in East Asia. Impatiens textori has considerably higher genetic diversity than I. hypophylla . For example, mean values of genetic diversity within populations (He) of I. textori and I. hypophylla were 0.206 and 0.102, respectively. The high outcrossing rate (mean 1 = 0.89), abundance and widespread distribution, and large population size may be factors responsible for the maintenance of high levels of genetic variation in I. textori . Analyss of fixation indices indicated that considerable population substructuring within populations of I. hypophylla , partly resulting from inbreeding and/or genetic drift in small populations coupled with limited gene flow (mean F_IS= 0.699). However, the degree of population divergence observed in I. hypophylla (mean G_ST= 0.059) was considerably lower than that in other plants with similar life history traits. It is supposed that similar selection at populations of I. hypophylla examined might maintain similar gene frequencies in isolated populations.     

长角凤仙花（凤仙花科）的花器官发生和发育

以不同发育时期的长角凤仙花Impatiens longicornuta Y.L.Chen（凤仙花科Balsaminaceae）为材料,利用扫描电镜技术观察了其花器官的分化及其发育过程。长角凤仙花为两侧对称花,具2枚侧生萼片,唇瓣囊状,旗瓣具鸡冠状突起,雄蕊5枚,子房上位,5心皮5室。其花器官分化顺序为向心式,萼片—花瓣—雄蕊—雌蕊原基。2枚侧生萼片先发生,然后近轴萼片（即唇瓣）原基和2枚前外侧萼片原基近同时发生;但是这3枚萼片原基的发育不同步,远轴的2枚前外侧萼片原基的发育渐渐滞后,然后停止发育,最后渐渐为周围组织所吸收,直至消失不见。花瓣原基中,旗瓣原基最先发生,4个侧生花瓣原基相继成对发生,且之后在基部成对愈合形成翼瓣;5枚雄蕊原基几乎同时发生,5个心皮原基轮状同时发生。本文结果支持凤仙花属植物为5基数的花,并进一步证实了唇瓣的萼片来源;此外,研究结果表明花器官早期发育资料对植物系统与进化研究具有重要参考价值。     

Four New Species of Impatiens (Balsaminaceae) from China

In the paper four new species of the genus Impatiens (Balsaminaceae) are described from China. They are Impatiens fenghwaiana Y. L. Chen, I. wuyuanensis Y. L. Chen,I. jinggangensis Y. L. Ghen, and 1. hunanensis Y. L. Chen.     

海南凤仙花与龙州凤仙花花的形态补充描述

凤仙花属植物的花部形态历来是该属的属下划分及种间界定的关键性状,基于大量的有花标本及野外实地调查,并参照了模式标本对海南凤仙花和龙州凤仙花的花部形态进行了补充描述。     

Testing for stress-dependent inbreeding depression in Impatiens capensis (Balsaminaceae)

The relevance of inbreeding depression to the persistence of plant populations can depend upon whether stress magnifies inbreeding depression for fitness-related traits. To examine whether drought stress exacerbates inbreeding depression in gas exchange traits and biomass, we grew selfed and outcrossed progeny of inbred lines from two populations of Impatiens capensis in a greenhouse experiment under water-limited and moist soil conditions. Drought stress did not magnify the degree of inbreeding depression for any of the traits measured. In fact, in one population there was a trend for stronger inbreeding depression under well-watered, benign conditions. Furthermore, significant inbreeding depression for carbon assimilation rate and stomatal conductance was only detected in the lines from one population. In contrast, inbreeding depression for biomass was detected within both populations and differed among lines. Drought stress exerted significant selection on physiological traits, favoring increased carbon assimilation rates and decreased stomatal conductance in drought-stressed plants. Patterns of selection did not differ between inbred and outcrossed plants but did differ marginally between populations. Thus, estimates of selection were not biased by the mixed mating system per se, but may be biased by combining individuals from populations with different histories of selection and inbreeding.     

Delayed autonomous self-pollination in Hibiscus laevis (Malvaceae)

Delayed autonomous self-pollination allows outcrossing to occur while also ensuring that seeds are produced in the absence of pollen vectors. We investigated variation in the efficacy of this pollination mechanism in populations of Hibiscus laevis. Recurvature of stylar branches occurred after 1 d of anthesis, and in plants from Ohio, Illinois, Arkansas, and Oklahoma this behavior resulted in autonomous selfing (surprisingly, stylar movement was facultative in that it did not take place when the stigmas were already pollinated). In contrast to these more northern populations, the distance between anthers and stigmas was too great to allow autonomous selfing in plants from Texas, Mississippi, and Louisiana. Greenhouse studies of plants from Ohio demonstrated that autonomous selfing resulted in an average of 29.5 seeds per flower, as compared to 59.9 seeds per flower from hand-pollination of stigmas with self pollen. In an assessment of the possible significance of this selfing mode, emasculated flowers did not set significantly fewer seeds in a natural stand in Ohio, suggesting that few seeds resulted from autonomous selfing at that site. Modest inbreeding depression was detected at this population. Our results suggest that delayed autonomous selfing is more common in northern populations, where it may facilitate population establishment and persistence at times when pollinators are scarce.     

The efficiency of insect pollination of the neophyte Impatiens glandulifera (Balsaminaceae)

To comprehend the reproductional success of Impatiens glandulifera in Central Europe, the pollination system was analyzed under field conditions. Impatiens glandulifera attracts a lot of visitors because of its abundant amount of nectar with its high sugar content. The range of visitors, their composition and their changes during the blooming time and their pollen loading capacity were investigated as well as plant and flower morphology, the nectar production, the quantity and viability of pollen, the stigmatic pollen load, the pollen/ovule ratio and the seed production of the neophyte. The visitors, which in Central Europe almost exclusively utilize the nectar, are passively covered with pollen because of the morphology and the mobile suspension of the flowers. Consequently the next landing results in a pollination of I. glandulifera during the female blooming phase. Although the germination rate of the pollen deposited on the stigma is very low, a 100% seed set is readily obtained during the initial visit to a flower in female stage.     

Hybridization and regional variation in Pacific Northwestern Impatiens (Balsaminaceae)

Variation patterns ofImpatiens species in the Pacific Northwest are discussed. Much of the variability there is attributed to widespread interspecific hybridization. Taxonomy of the genus is briefly reviewed, and it is suggested that four species should be recognized for the region. An account of natural hybridization betweenI. capensis andI. ecalcarata is given in detail, and it is postulated that the effects of this hybridization may extend beyond the immediate area in which it is detectable on the basis of morphological patterns. It is suggested thatI. aurella may be of hybrid origin since it combines characters of two species in the Pacific Northwest. Although natural hybridization is occurring at the present time, geographical distribution of variant populations of the species suggests that extensive hybridization may have been initiated in late Pleistocene as a result of the ecological disturbances prevalent at that time.     

Kin recognition: Competition and cooperation in Impatiens (Balsaminaceae)

The ability to recognize kin is an important element in social behavior and can lead to the evolution of altruism. Recently, it has been shown that plants are capable of kin recognition through root interactions. Here we tested for kin recognition in a North American species of Impatiens that has a high opportunity of growing with kin and responds strongly to aboveground competition. We measured how the plants responded to the aboveground light quality cues of competition and to the presence of root neighbors and determined whether the responses depended on whether the neighbors were siblings or strangers. The study families were identified by DNA sequencing as members of the same species, provisionally identified as Impatiens pallida (hereafter I. cf. pallida). We found that I. cf. pallida plants were capable of kin recognition, but only in the presence of another plant's roots. Several traits responded to relatedness in shared pots, including increased leaf to root allocation with strangers and increased stem elongation and branchiness in response to kin, potentially indicating both increased competition toward strangers and reduced interference (cooperation) toward kin. Impatiens cf. pallida responded to both competition cues simultaneously, with the responses to the aboveground competition cue dependent on the presence of the belowground competition cue.     

华凤仙花部特征和传粉系统研究

研究了华凤仙（Impatiens chinensis L.）的花部特征和传粉系统,结果表明华凤仙单花期为2.3 d,雄性期是雌性期的6.8倍。花蜜量和花蜜可溶性糖含量分别为0.56 μL和28.2%。开花当天柱头即具有可授性,花粉活力在花朵刚开放时最高（95%）,至花朵凋谢之前仍保持较高的活力（55%）。花粉胚珠比（5 730.3±2 941.1）和套袋实验的结果表明华凤仙属于自交亲和但以异交为主的繁育系统,传粉过程需要传粉者。蜜蜂、熊蜂、蝶类和几种芦蜂是华凤仙的主要访花者,最有效的传粉者为三条熊蜂。考氏无垫蜂作为盗蜜者,可能是除正常访花者之外,对华凤仙的有性繁殖影响最大的生物因素。     

Development of thirteen microsatellite loci for Impatiens lateristachys (Balsaminaceae)

Impatiens lateristachys is a local endemic species of Emei Mountain in southwestern China. In order to study the speciation mode of this species and its closely relatives, it is important to characterize their genetic diversity and differentiation. Here, thirteen polymorphic microsatellite markers have been developed for I. lateristachys. AC/TG microsatellite was enriched by combining biotin capture method. The average allele number of these markers was 3.1, ranging from 2 to 6. The observed (H _O) and expected (H _E) heterozygosities were 0.15−0.55 and 0.21–0.70, respectively. All the 13 markers could be applied in the other four congeneric species.     

海南凤仙花不同海拔种群的传粉生物学

凤仙花属(Impatiens)植物具有极为广泛的多样性和类型各异的特化传粉者, 被誉为“双子叶的兰花”, 受到众多传粉生物学家的关注。本文以海南特有种海南凤仙花(Impatiens hainanensis)为研究对象, 对3个不同海拔梯度的种群进行了开花生物学特性和开花物候、花器官结构、花粉活力和柱头活性、传粉者种类和访花行为及繁育系统的比较研究。结果表明: 单花花期4.10 ± 0.46 d, 雄性期和雌性期分别约为3.15 ± 0.24 d和0.95 ± 0.36 d; 种群开花峰期在8月初, 高海拔种群的花期高峰相对滞后。低、中海拔种群花粉活力呈现先升高后下降的趋势, 以开花第2 d花粉活力最高; 高海拔种群花粉活力随开花时间推移逐渐下降; 柱头活性随开花时间的推移而增强, 高海拔种群开花各天次均较低、中海拔种群低。主要传粉昆虫为黄黑无垫蜂(Amegilla leptocoma)和绿条无垫蜂(A. zonata), 低、中海拔种群以黄黑无垫蜂为主, 高海拔种群以绿条无垫蜂为主。未观察到自动自花授粉和无融合生殖现象, 人工授粉能明显增加坐果率(75-90%), 自然坐果率在高海拔种群相对较低(40-60%), 说明存在较强的传粉者限制。海南凤仙花的保护需要同时关注其有效传粉者的保护, 促进有效传粉昆虫在不同海拔种群之间的往来, 保证种群间的花粉流与种子流, 维持海南凤仙花的种群遗传多样性与有效种群大小。     

Maternal effects of drought stress and inbreeding in Impatiens capensis (Balsaminaceae)

Maternal effects can have substantial impacts on plant fitness and plant populations. Stressful environmental conditions can cause a maternal plant to inadequately provision its progeny, resulting in poor seedling growth, low reproductive success, and decreased competitive ability. Maternal effects consist of environmental and genetic load components, but the interactions between these two components have rarely been considered. To determine the effects of maternal drought stress and maternal inbreeding on progeny biomass (a fitness correlate) and physiological responses to drought stress, we conducted a greenhouse experiment with genetic lines from two populations (mesic site vs. dry site) of the herbaceous annual Impatiens capensis (Balsaminaceae). Seeds were collected from cleistogamous flowers of inbred or outcrossed maternal plants that were subject to either a drought or control treatment. These seeds were grown into juvenile plants that were also subject to either a drought stress or a control treatment. Plants from the mesic site had significantly reduced biomass from maternal drought stress, while plants from the dry site maintained biomass despite adverse maternal environmental conditions. Juvenile plants of both populations had reduced biomass only as a result of maternal inbreeding. Interestingly, inbreeding depression was more apparent when maternal environmental conditions were benign.     

Foraging behavior affects pollen removal and deposition in Impatiens capensis (Balsaminaceae)

Flowers of most plant species are visited by a variety of animals. Some of these visitors are effective pollinators while others remove resources without transferring pollen. Studies comparing the effectiveness of different visitors as pollinators often compare taxa without considering variation in behavior within a taxon. Wilson and Thomson (Ecology 72: 1503-1507, 1991) documented the effects of honey bees and bumble bees on the pollination dynamics of Impatiens capensis. They found that pollen-collecting honey bees removed large numbers of pollen grains from anthers but deposited little of it on stigmas; bumble bees, which sought nectar, removed less pollen but deposited more of it on stigmas. It is unclear whether the low pollen transfer efficiencies of honey bees are explained by their morphology or by their pollen-collecting behavior. We repeated the work of Wilson and Thomson at a site where honey bees were foraging for nectar, not pollen. We measured the quantity of pollen remaining in anthers, the number of pollen grains deposited on stigmas, and seed production after single visits by honey bees and bumble bees. The differences between the taxa disappeared when they were foraging in a similar manner. Our results clearly demonstrate the importance of foraging behavior on the pollination effectiveness of floral visitors.     

Impatiens maculifera sp. nov. (Balsaminaceae) Yunnan,China

Impatiens (Balsaminaceae) is a notoriously difficult genus to study for reasons that include small, remote and endemic areas of occurrence, and difficulty in obtaining well‐dried herbarium specimens and separating and reconstructing floral parts from available dried specimens. Impatiens has also been notoriously difficult when it comes to phylogenetic resolution at the infrageneric level, but a new system with two subgenera (Impatiens and Clavicarpa) was recently proposed by Yu et al. and it has made it easier to classify newly discovered species. Impatiens maculifera, a new species of Balsaminaceae from Malipo County, Yunnan, China described here, is similar to I. parvisepala S.X. Yu & Y.T. Hou in having racemose inflorescences, four lateral sepals, clavate capsules and ellipsoid seeds, but differs in having leaves narrowly oval or with elliptic–lanceolate blades (versus obovate or obovate–lanceolate), obvious petioles (versus leaves subsessile), racemose inflorescence with 8–12 flowers (versus 6–8 flowers), flowers greenish yellow (versus yellow), lateral united petals and lower sepal with dense red spots (versus red spots absent), and dorsal petal with apparent stalk at base (versus unapparent stalk). Morphological characters and molecular phylogenetic analyses of DNA sequences from both nuclear ribosomal and plastid genes show that the new species differs distinctively from I. parvisepala. Furthermore, I. maculifera is distinguished from other Impatiens species based on morphological, micromorphological and palynological evidence, and molecular data.     

Impatiens pyrorhiza sp. nov. (Balsaminaceae) from east Himalaya

Impatiens pyrorhiza sp. nov. from the Tawang and West Kameng districts in northwest Arunachal Pradesh, India, is described and compared with related species. It is similar to I. gamblei from east Nepal and northwest Darjeeling, but differs in for example: a) rhizome extended, orange, much‐branched, b) leaves with numerous teeth, c) petiole base with prominent glands, d) raceme supported by subtending leaf and e) lateral sepals without an apical divergent extended callosity.