雌雄异熟植物露蕊乌头开花时间对雌雄功能期及表型性别的影响

开花时间决定了植物雌雄功能的交配机会, 最终影响繁殖成功。交配环境假说认为雌雄异熟植物开花时间的差异能引起植物表型性别的变异, 改变种群内的交配环境, 影响植物对雌雄功能的最佳性分配。为了研究开花时间对雌雄异熟植物的雌雄性别时期及表型性别的影响, 本文以毛茛科雄性先熟植物露蕊乌头(Aconitum gymnandrum)为实验材料, 记录了雄性和雌性功能期, 分析了植株开花时间、花的雌雄功能期和表型性别的关系。结果表明: 在植物同一花序内, 较晚开放的花有更长的雄性期和更短的雌性期, 性分配在时间上偏雄。雌雄功能期在时间上的相对分配随植物开花时间的变化表现出相似的趋势： 较晚开的花或较晚开花的个体, 花的雄性功能期相对于雌性功能期更长, 在时间上更偏向雄性功能。而且, 开花时间的差异影响种群内花的性比和植物个体的表型性别动态。随着开花时间由早到晚的变化, 种群内早期以雄花为主,末期以雌花为主, 种群内性别环境由偏雄向偏雌变化, 因此植株个体的平均表型性别则从偏雌转向偏雄。本文结果支持交配环境假说, 雄性先熟的露蕊乌头开花早期, 种群内花的性别比偏雄, 种群表型性别环境偏雄, 因而植物个体平均表型性别偏雌, 性别分配(即时间分配)偏向雌性功能, 而晚开花个体的平均性别偏雄, 更偏向雄性功能的分配。     

优质荔枝新品种''草莓荔''开花与落果特性观察

从植株群体和单穗调查分析‘草莓荔’雌雄花开花特性,结果表明,‘草莓荔’雌雄花交替开放的顺序主要是雌→雄,即前期开雌花,后期开雄花;但也有少量交替开放顺序是雄→雌→雄,即先开雄花,中期开雌花后再开雄花。植株群体的花期和雌雄花开放相遇时间比单穗长,更有利于传粉受精。落果从花后15d开始出现,并在花后18～25d出现第一次落果高峰,落果最多,开花后51～58d为第二次落果高峰。     

优质荔枝新品种‘草莓荔’开花与落果特性观察

从植株群体和单穗调查分析‘草莓荔’雌雄花开花特性,结果表明,‘草莓荔’雌雄花交替开放的顺序主要是雌→雄,即前期开雌花,后期开雄花;但也有少量交替开放顺序是雄→雌→雄,即先开雄花,中期开雌花后再开雄花。植株群体的花期和雌雄花开放相遇时间比单穗长,更有利于传粉受精。落果从花后15d开始出现,并在花后18～25d出现第一次落果高峰,落果最多,开花后51～58d为第二次落果高峰。     

元宝枫花芽分化及花开放过程研究

为探明元宝枫花性别、交配系统及花芽分化过程,通过石蜡切片和连续解剖观察,对元宝枫花开放及花芽分化过程进行了研究。结果表明：(1)元宝枫花性别可分为雌能花和雄能花两种,雌能花花药不开裂,只表现雌性功能,而雄能花分Ⅰ型和Ⅱ型两种,花药均可产生成熟花粉并开裂散粉,只表现雄性功能。(2)元宝枫是较为少见的二重雌雄异型异熟树种,且交配系统有雄先型和雌先型两种;雄先型小花开放顺序为：雄能花→雌能花→雄能花,雌先型小花开放顺序为雌能花→雄能花→雄能花。(3)元宝枫花芽的形态分化期开始于7月上旬,花序和小花分化期为8月上旬,雄蕊和雌蕊分化开始于8月下旬,雌配子体发育晚于雄配子体。(4)元宝枫花性别分化的关键期为第二年的3月下旬至4月上旬。     

基于形态学及生理学的黄檗性别差异研究

为了探究黄檗雌雄植株的形态学及生理学差异,对黄檗成树雌雄植株的形态学、叶片抗氧化防御酶活性及内源激素含量等指标开展对比研究,结果表明：黄檗雌雄植株形态存在显著差异,雌株叶宽比雄株大1.66±0.148 cm（P<0.05）、雌株枝间夹角比雄株大17.5±1.21°（P<0.001）;雌雄株叶片颜色存在差异,但叶绿素含量差异不显著;黄檗雌雄株超氧化物歧化酶（SOD）、过氧化物酶（POD）、过氧化氢酶活性（CAT）酶活性存在显著差异,CAT酶活性在7月最高,雌株含量大雄株;POD酶活性6月含量最高,雄株大于雌株;SOD酶活性8月最高,雄株大于雌株。内源激素赤霉素（GA3）含量雄株大于雌株;吲哚乙酸（IAA）含量为雌株大于雄株;玉米素（ZA）含量为雄株大于雌株,差异显著（P<0.05）;内源脱落酸（ABA）性别间无显著差异。黄檗雌雄植株在形态及生理学等指标上存在一定差异,说明黄檗雌雄植株为维持自身生长需要,在形态和生理方面都做出不同调整以适应生存环境。     

棉铃虫温度依赖的生长发育及雌性先熟现象

在昆虫中,雄虫先于雌虫羽化,称为雄虫先熟现象,而雌虫先于雄虫羽化,称为雌虫先熟现象。前者最为普遍,后者报道较少。本试验在20、22和25℃下对棉铃虫Helicoverpa armigera（Hilbner）的生长发育进行了详细的研究。结果显示发育历期随温度的升高而缩短,雌雄间发育历期有着显著的差异,各个温度下雌虫均比雄虫先羽化,表明棉铃虫属于雌性先熟种类,而这种差异主要是由于雌虫的蛹期较短所致。幼虫发育历期与蛹重呈显著的正相关性。     

脓疮草复合体的生物学特性研究

通过野外和移栽实验观测,对脓疮草复合体（Ｐａｎｚｅｒｉａｎ ｌａｎａｔａ ｃｏｍｐｌｅｘ）居群进行了生物学特性研究。结果表明：浓疮草复合体居群在生活史方面无明显差异,移栽的１０个复合体居群生活史十分 相似：脓疮草复合本的花量大,花期可达３个月左右,花为雌雄异熟,以异花授粉为主,自交（同株异花授粉）亲合;叶裂程度、花萼裂片宽窄等性状,在天然居群中是连续变化的,而且移栽后可逆性较大,性状不稳定。     

青杨雌雄群体沿海拔梯度的分布特征

以小五台山天然青杨种群为试验对象,通过在其分布海拔范围内(1400-1700 m)设置样方的方式研究青杨雌雄群体的平均胸径、密度、性比、大小级结构和空间分布的差异.结果显示:雌株群体的平均胸径在整个海拔梯度上无显著差异,而海拔1700 m处的群体密度显著低于其它海拔;雄株群体的平均胸径在海拔1700 m最大,显著高于其它海拔,而群体密度在各海拔梯度间无显著差异.在海拔1600 m,雌雄群体的平均胸径最接近,同时密度也最接近.从整个海拔范围来看,青杨雌雄个体的比例(雄/雌)为0.80∶1,性比不偏离1∶1(x2=2.94,P＞0.05).但在不同海拔梯度上性比有所不同,低海拔性比为0.44,显著偏雌(X21400=5.91,P＜0.05),而高海拔性比为2.55,显著偏雄(x21700=6.56,P＜0.05);随着海拔接近1600 m,性比逐渐趋于1:1(x21600=0,P＞0.05).除海拔1700 m为下降型外,青杨雌雄群体的大小级结构在其它海拔主要表现为稳定型.而最稳定群体结构的分布海拔在雌雄群体间不同.在海拔1400 m,雌株群体中I级和Ⅱ级植株占群体比例最大;雄株群体I级和Ⅱ级植株占群体比例最大的海拔是1600 m.群体空间分布以聚集分布为主,彼此间无明显差异;但群体间聚集强度的变化在海拔梯度上各有不同,主要表现为雌株群体的PAI随海拔的升高逐渐增加,而雄株群体的PAI则随海拔的升高呈现出先增加后减少的趋势.研究结果表明了青杨雌雄群体的分布特征沿海拔梯度的变化明显不同,中等海拔区域可能为青杨种群的最适繁衍区.     

海滨蝇子草功能性雌全同株性系统的初步研究

雌全同株是指植株同时具有两性花和雌花性系统,目前国内外相关研究多集中于菊科（Asteraceae）植物。海滨蝇子草（Silene uniflora Roth.）单株具有2种花朵类型,分别是可育两性花和功能上为雌花的雄性不育两性花,属于功能性雌全同株植物。可育两性花具有雄性先熟和雌雄异位的性别分离特征;其繁育系统主要为异交,部分自交,花内自交坐果率为31.71%。功能性雌花的雄蕊和花粉畸形,花粉量少。在初花期和末花期,功能性雌花较可育两性花所占比例更大。海滨蝇子草的功能性雌全同株性系统,在进化上可能具有降低自交、增加异交率,避免雌雄功能干扰和提高雌雄功能资源分配等适应性机制。     

Temperature-dependent development and protogyny in Helicoverpa armigera

在昆虫中,雄虫先于雌虫羽化,称为雄虫先熟现象,而雌虫先于雄虫羽化,称为雌虫先熟现象.前者最为普遍,后者报道较少.本试验在20、22和25℃下对棉铃虫Helicoverpa armigera(Hübner)的生长发育进行了详细的研究.结果显示发育历期随温度的升高而缩短,雌雄间发育历期有着显著的差异,各个温度下雌虫均比雄虫先羽化,表明棉铃虫属于雌性先熟种类,而这种差异主要是由于雌虫的蛹期较短所致.幼虫发育历期与蛹重呈显著的正相关性.     

Flowering phenology and wind-pollination efficacy of heterodichogamous Juglans mandshurica (Juglandaceae)

BACKGROUND AND AIMS: Heterodichogamy differs from normal dichogamy, in that it involves two mating types (protogyny and protandry) that occur at a 1 : 1 ratio in a population. Flowering phases of the two mating types are synchronized and reciprocal, which was considered to ensure between-type outcrossing. This study aims to quantify the flowering pattern and pollination efficacy in Juglans mandshurica, a wind-pollinated heterodichogamous tree. METHODS: The pattern of flowering phenology was monitored within individual trees and pollen traps were used to measure air-borne pollen loads during the spring in 2003 and 2004. Pollen longevity was determined by staining technique. Also a pollen supplementation experiment was performed in 2004 to assess pollen limitation of fruit production. KEY RESULTS: There was no overlap between sexual functions within individual trees. Flowering periods of the two mating types were reciprocal and synchronous in both 2003 and 2004. Air-borne pollen loads were large, and protogynous and protandrous individuals each produced a high pollination peak, consistent with the two blooming periods. Maximum pollen longevity was about 4 h for protandrous individuals, and 3 h for protogynous individuals. Pollen supplementation did not increase fruit production in either protogynous or protandrous individuals. CONCLUSIONS: Heterodichogamous flowering in Juglans mandshurica effectively avoids selfing, promotes between-type outcrossing, and leads to efficient pollination in a natural population.     

The relationship between phenology of pistillate flower organogenesis and mode of heterodichogamy in Juglans regia L. (Juglandaceae)

 We investigated the degree of organogenesis completed at the end of the growing season in pistillate flowers of heterodichogamous Juglans regia, English or Persian walnut. Terminal buds from paired cultivars, one each protandrous and protogynous, chosen to represent early, midseason and late leafing walnuts, were examined by scanning electron microscopy. Results indicate that pistillate floral primordia in protandrous individuals had not progressed beyond involucre initiation during the season prior to bloom. In protogynous individuals, floral differentiation had progressed to the initiation of perianth primordia. These observations are compared with an earlier report on staminate flower differentiation in the same cultivars where a comparable, but opposite, relationship exists. We conclude that the degree of differentiation in both staminate and pistillate flowers that must be completed between the time growth resumes in the spring and anthesis is a developmental determinant of the mode of heterodichogamy in walnut. Received: 15 June 1996 / Revision accepted: 25 October 1996     

HETERODICHOGAMY IN GRAYIA BRANDEGEI (CHENOPODIACEAE): REPORT FROM A NEW FAMILY

The reproductive biology of Grayia brandegei was examined. Although monoecious, Grayia brandegei exhibits a phenotypic dimorphism of protogynous and protandrous mating types known as heterodichogamy. For individual plants, the temporal separation of staminate and pistillate flowering phases appears to be complete. No self-fertilization is possible in protandrous plants, but may be possible in protogynous plants provided pistillate flowers remain unfertilized. Flowering phases of protogynous and protandrous mating types are synchronized and reciprocal, ensuring cross-fertilization between mating types. Less than 15% overlap of sexual functions occurred between plants of the same mating type. Protogynous and protandrous plants were randomly dispersed in the environment and in relation to each other. Mating type frequencies did not differ significantly from 1:1. We discuss the possibility of a heterodichogamous pathway to dioecy.     

Gender specialization and factors affecting fruit set of the wind-pollinated heterodichogamous Juglans regia

Gender expression, flowering phenology, reproductive performance and factors affecting fruit set (i.e., flowering synchrony, size and distance to the nearest pollen donor) were investigated in a cultivated population of a wind-pollinated self-compatible heterodichogamous Juglans regia (Juglandaceae). Four flowering morphs, (i.e., protandrous, protogynous, male and female) were observed. The sexual functions of the protandrous and protogynous morphs were almost synchronous; however, they were not reciprocal, and the separation of male and female flowering within most monoecious individuals was not complete. Thus, within-morph mating and geitonogamous pollination may be common. The ratio of protandrous versus protogynous morphs was biased towards the protandrous morph, but the fruit set did not differ between the morphs, suggesting that the fruit set of the protandrous morph could be partly compensated by within-morph pollination. The ratio of the female flower number or fruit number to the total male catkin length was higher in the protogynous morph than in the protandrous morph and did not vary with plant size, suggesting that gender variation was not size dependent and that the sexual function of protandrous morphs was more male biased. Fruit set depended on plant size only for protandrous morphs. The fruit set of individual plants decreased with increasing distance to the nearest pollen donor regardless of morph, possibly because of pollen limitation. The fruit set of individual plants increased with flowering synchrony, indicating that flowering synchrony could affect reproductive success.     

Dichogamy in angiosperms

We obtained information on dichogamy and other aspects of the biology of over 4200 species of angiosperms from several hundred published and unpublished sources. We used this information to describe patterns of occurrence of dichogamy and to test specific hypotheses relating dichogamy to other characteristics of plants or their environments. Protandry was more common than protogyny at the intrafloral level, but the reverse was true at the interfloral level. Patterns of dichogamy varied significantly among major taxa, with protogyny more common among monocotyledons and primitive dicotyledons, and protandry expecially common in the Asteridae. Arctic species tended to be less dichogamous and more protogynous than temperate and tropical species. Aquatic and alpine species were especially protogynous. Patterns of dichogamy varied among sexual systems, with gynomonoecious and gynodioecious species especially protandrous, and monoecious species highly protogynous. Autogamous and self-compatible species were disproportionately protogynous. Flowers of intraflorally dichogamous species were slightly larger than those of adichogamous species, owing to the presence of many autogamous species in the latter group. Species with interfloral protogyny bore much smaller flowers than did species with interfloral protandry. Early-blooming species in north-temperate and polar regions were disproportionately protogynous. Sexual structures that abscised, shriveled or moved after completion of their function tended to be presented first, and those that facilitated the other sexual function were presented second. A negative association existed between type of intrafloral and interfloral dichogamy in diclinous species. Most animal-pollinated flowers were protandrous, except beetle-pollinated and refuge and trap blossoms. Wind pollination was markedly associated with protogyny. Vertical inflorescences visited by upwardly-moving vectors were protandrous.     

Temporal floral sex allocation in protogynous Aquilegia yabeana contrasts with protandrous species: support for the mating environment hypothesis

We tested one of the predictions of Brunet and Charlesworth (1995) that relative floral sex allocation will vary temporally with the mating environment and that the form of dichogamy (protandry vs. protogyny) will select for the pattern of variation in male versus female resource allocation. In many hermaphroditic plant species, allocation to female function (ovule number) decreases from early to late flowers within inflorescences as a result of resource limitation or ontogenetic changes. This pattern may obscure the effects of the mating environment and dichogamy on selection for allocation patterns in protandrous species (male allocation increases regardless). By examining a protogynous species the alternative pattern of temporal variation in resource allocation is predicted, namely that allocation to male function should decrease (or female allocation increase) throughout the flowering sequence. This pattern was observed in protogynous Aquilegia yabeana (Ranunculaceae), in which ovule number per flower remained constant whereas pollen number decreased in sequentially blooming flowers. These observations support the temporal sex allocation hypothesis of Brunet and Charlesworth (1995).     

Barriers to interspecific hybridization between Juglans nigra L. and J. regia L species

Juglans nigra and Juglans regia are phylogenetically divergent species. Despite the economic interest in Juglans?×?intermedia (J. nigra?×? J. regia), walnut hybridization is rare under natural conditions and still difficult using controlled pollination. Here, we evaluated some reproductive mechanisms that may prevent successful natural hybridization. The study of flowering phenology of 11 J. nigra and 50 J. regia trees growing in a plantation provided information regarding the opportunity for interspecific crosses. Variation in flower size, pollen quality of putative donors, and variation in seed yield and rate of hybrid production among putative maternal trees were examined. DNA fingerprinting and parentage analyses based on nine microsatellites permitted the identification of hybrids and hybridogenic parent. Our data indicated that overlap occurred between the staminate flowering of protogynous J. regia and the beginning of pistillate flowering of protogynous J. nigra. Differences in floral size were computed between walnut species. Only three hybrids among 422 offspring of eleven J. nigra progenies were identified. Interspecific hybridization involving pollination of one early-flowering-protogynous J. nigra by three protogynous J. regia trees was detected. The correct development of J. regia male gametophytes, high pollen viability (86.5 %), and germination (57.6 %) ruled out the possibility that low pollen quality contributed to depressed hybrid production. Our findings indicated that these two species tended to remain reproductively isolated. The substantial disjunction in flowering time and additional prezygotic barriers such as differences in floral size and conspecific pollen advance may affect interspecific gene flow between J. regia and J. nigra.     

Resource partitioning to male and female flowers of Spinacia oleracea L. in relation to whole-plant monocarpic senescence

Male plants of spinach (Spinacea oleracea L.) senesce following flowering. It has been suggested that nutrient drain by male flowers is insufficient to trigger senescence. The partitioning of radiolabelled photosynthate between vegetative and reproductive tissue was compared in male (staminate) versus female (pistillate) plants. After the start of flowering staminate plants senesce 3 weeks earlier than pistillate plants. Soon after the start of flowering, staminate plants allocated several times as much photosynthate to flowering structures as did pistillate plants. The buds of staminate flowers with developing pollen had the greatest draw of photosynthate. When the staminate plants begin to show senescence 68% of fixed C was allocated to the staminate reproductive structures. In the pistillate plants, export to the developing fruits and young flowers remained near 10% until mid-reproductive development, when it increased to 40%, declining to 27% as the plants started to senesce. These differences were also present on a sink-mass corrected basis. Flowers on staminate spinach plants develop faster than pistillate flowers and have a greater draw of photosynthate than do pistillate flowers and fruits, although for a shorter period. Pistillate plants also produce more leaf area within the inflorescence to sustain the developing fruits. The (14)C in the staminate flowers declined due to respiration, especially during pollen maturation; no such loss occurred in pistillate reproductive structures. The partitioning to the reproductive structures correlates with the greater production of floral versus vegetative tissue in staminate plants and their more rapid senescence. As at senescence the leaves still had adequate carbohydrate, the resources are clearly phloem-transported compounds other than carbohydrates. The extent of the resource redistribution to reproductive structures and away from the development of new vegetative sinks, starting very early in the reproductive phase, is sufficient to account for the triggering of senescence in the rest of the plant.     

Flowering and fruiting phenology of Normanbya normanbyi (W. Hill) L. H. Bailey (Arecaceae), a palm endemic to the lowland tropical rainforest of north‐eastern Australia

Abstract Normanbya normanbyi (W. Hill) L. H. Bailey (Arecaceae) is a monoecious, arborescent palm with a very small distribution area within the Daintree rainforest in north‐eastern Australia. Our 2‐year study was focused on the reproductive phenology at the individual and population level. At the population level flowering peaked in the dry season, whereas fruiting was confined to the wet season. Each palm can bear up to three inflorescences/infructescences at the same time. Flowering of each inflorescence is separated from each other by a couple of weeks. A single inflorescence consists of about 1900 staminate and 800 pistillate flowers. The flowering of N. normanbyi is protandrous with a staminate phase lasting 40 days and a pistillate phase of approximately 2 weeks. Between both phases is a non‐flowering phase of about 9 days. Fruit ripening takes 21 weeks, with an average of about 280 ripe fruit per tree. Comparison of three study plots revealed a moderate synchrony of flowering and fruiting initiation in this species of palm. The male phase of flowering shows a higher degree of synchrony than the female phase at the population level. Seasonal regularity of flowering and fruiting peaks appears to be predictable. The general flowering and fruiting phenology of N. normanbyi follows a subannual pattern with a strong tendency towards a continual pattern.     

Pollen movement to flowering canopies of pistillate individuals of three rain forest tree species in tropical Australia

This study measured the quantities of effective pollen vectors and their pollen loads arriving at the canopies of dioecious tropical rain forest trees in north-east Queensland. Population flowering synchrony, effective pollinator populations and pollen loads transferred between staminate and pistillate trees were compared among three insect-pollinated tree species. All three were visited by a wide range of insects, 75% of which (mostly 3–6 mm long) carried conspecific pollen. Fewer than 8% of individual insects were found to be carrying single-species pollen exclusively and none could be described as specialist pollen foragers. The introduced honeybee carried greater quantities of pollen than any native species but was not necessarily a reliable pollinator. The brief flowering periods in Neolitsea dealbata (3–4 weeks) and Litsea leefeana (4–5 weeks) populations were synchronized among individuals. Flowering in the Diospyros pentamera population extended over 15 weeks and most individuals were in flower for most of this period. Staminate trees began flowering earlier, produced more flowers and attracted relatively more insects than did pistillate trees, suggesting a density-dependent response of pollinators to flowering performance. Pollen was trapped in greater quantities on insects at staminate trees than at pistillate trees. Insect numbers increased at peak flowering periods and Diptera were the most abundant flower visitors. Anthophilous Coleoptera were more numerous at staminate than at pistillate trees in all three tree species populations. Larger quantities of pollen were mobilized during peak flowering times although the greatest quantities were transferred to pistillate canopies towards the end of the population flowering periods. Diptera carried pollen more often to pistillate N. dealbata and L. leefeana trees than did other groups whereas Coleoptera carried pollen more often to pistillate D. pentamera trees. The two contrasting flowering performances in the three tree species are discussed with reference to mechanisms that facilitate pollen transfer between staminate and pistillate trees.