舞花姜属3种植物繁殖策略比较

选择自然分布于西双版纳地区的毛舞花姜(GlobbabartheiGagnepain)、双翅舞花姜(G.schomburgkiiJ.D.Hooker)和澜沧舞花姜(G.lancangensisY.Y.Qian)为实验材料,比较了3种植物的有性和无性繁殖特性。毛舞花姜和双翅舞花姜在自然状况下开花不结实,人工自交与异交、人工去除珠芽均不能使其恢复有性繁殖能力;花粉败育(毛舞花姜)或花粉活力低(双翅舞花姜,花粉萌发率<12%)可能是这2种植物不能结实的主要原因。毛舞花姜和双翅舞花姜均以珠芽为主要繁殖体,一年生植株每花序产珠芽分别为16.46±3.56(平均值±标准差,N=60)和14.25±3.63(平均值±标准差,N=153)。澜沧舞花姜具有雄花两性花同株的性表达特征,自然状态下以种子繁殖为主,结实率(种子/胚珠比)可达64.5%±12.1%(N=36),人工自交结实率显著低于异交结实率,有明显的自交不亲和现象;花序上产生少量珠芽[每花序产珠芽2.24±1.41(平均值±标准差,N=184)]。此外,3种舞花姜属植物还具有通过地下茎进行克隆繁殖的能力。3种舞花姜属植物的拜访昆虫均为蜂类,毛舞花姜的拜访昆虫只有排蜂(Megapisdorstata)。双翅舞花姜拜访昆虫主要为排蜂、黄绿彩带蜂(Nomiastrigata)及木蜂(Xylocopasp.);澜沧舞花姜的拜访昆虫主要为排蜂和黄绿彩带蜂。3种舞花姜属植物的花冠管长度存在     

雄全异株植物瘿椒树（省沽油科）的传粉生物学

从开花动态、传粉昆虫、花的形态结构、繁育系统、花粉活力和柱头可授性等方面研究了我国特有珍稀植物瘿椒树（Tapiscia sinensis Oliv.）的传粉生物学特性。瘿椒树是典型的雄全异株植物,两性花中含有功能性花粉,且自交亲和,但雄花花粉活力和萌发力是两性花的10倍以上。雄株和两性植株具有相同开花物候期,花期均为5月下旬至6月上旬,单花期为4-5天,雄花和两性花的5枚花药开裂的不同步性明显延长了散粉时间。两性花雌蕊先熟,柱头可授性较长。具有适应风媒和虫媒传粉的花部特征。传粉昆虫主要为蜜蜂科（Apidae）和食蚜蝇科（Syrphidae）昆虫,访花高峰期为8：30-10：30。维持瘿椒树雄全异株的可能机制是：雄株总体上增加了异交花粉的数量和质量;两性花的雄蕊为该物种提供了繁殖保障,同时为传粉者提供了报酬。     

雄全异株植物瘿椒树(省沽油科)的传粉生物学

从开花动态、传粉昆虫、花的形态结构、繁育系统、花粉活力和柱头可授性等方面研究了我国特有珍稀植物瘿椒树(Tapiscia sinensis Oliv.)的传粉生物学特性。瘿椒树是典型的雄全异株植物,两性花中含有功能性花粉,且自交亲和,但雄花花粉活力和萌发力是两性花的10倍以上。雄株和两性植株具有相同开花物候期,花期均为5月下旬至6月上旬,单花期为4-5天,雄花和两性花的5枚花药开裂的不同步性明显延长了散粉时间。两性花雌蕊先熟,柱头可授性较长。具有适应风媒和虫媒传粉的花部特征。传粉昆虫主要为蜜蜂科(Apidae)和食蚜蝇科(Syrphidae)昆虫,访花高峰期为8:30-10:30。维持瘿椒树雄全异株的可能机制是:雄株总体上增加了异交花粉的数量和质量;两性花的雄蕊为该物种提供了繁殖保障,同时为传粉者提供了报酬。     

雄全异株流苏树的花部特征及繁育系统研究

雄全异株是自然界罕见的繁育系统。通过野外观察和人工授粉实验对雄全异株植物流苏树传粉生物学特征及繁育系统进行研究。结果表明:流苏树的雄花与两性花的雄蕊发育过程基本一致,均能产生功能花粉粒。两性花的两个心皮原基愈合分化形成雌蕊,雄花的两个心皮原基愈合后形成一个空室并停止发育至整体退化。雌蕊先熟,柱头可授期长,花粉在花药开裂后具有活力,室温下,活力维持在10%以上约2周。流苏树靠风和昆虫(主要是蓟马和食蚜蝇)传粉。控制授粉30 d后,自然对照结实率为34.36%;两性花不存在无融合生殖现象,自交亲和,但自发自交的结实率仅10.70%;人工授粉下杂交结实率显著高于自交(同株异花);有性生殖受到传粉者限制;是混合交配系统。证实流苏树是木犀科又一功能性的雄全异株,其依靠雄株增加异交花粉的数量和质量,避免自交衰退;同时两性花的自交亲和保障生殖成功。流苏树雄花的雌蕊退化,从另一个角度证明木犀科的雄全异株是两性株向雌雄异株进化的过渡状态。     

白花鬼针草的繁殖特性及入侵性

白花鬼针草是一种菊科入侵植物。本研究从单个花序花期的观察、繁育系统、花粉胚珠比、自交亲和性、花粉活力、访花昆虫、种子萌发特性等方面对白花鬼针草的繁殖特性进行了分析,探讨了其与入侵性之间的关系。研究表明:白花鬼针草单个花序全部开放约5～7 d,约第9天全部凋落;花粉活力在1～8 d内维持在较高的水平(87.07%～93.41%);花粉胚珠比为2508,单个花序可自交亲和,亦可兼性异交,为混合交配系统,结实率高;边花(舌状花)的存在有利于结实率的提高;访花昆虫主要有粉蝶科、蛱蝶科、茧蜂科和蚁科的昆虫;对萌发率、萌发速率、萌发开始时间和萌发持续时间做因子及聚类分析,白花鬼针草种子的萌发特性可以表现为爆发型、过渡型、缓萌型、低萌型4种萌发类型,根据不同的环境条件选择不同的萌发类型。总之,白花鬼针草具有自交亲和,结实率高,结实量大,种子萌发率高、萌发速率快且具有不同的萌发策略和边花等繁殖特点,是其成功入侵的主要原因。     

野生翠雀传粉生物学特性研究

对从北京郊区引种的野生翠雀(Delphiniun grandiflorum L.)的开花习性进行了观察,对其花粉活力、柱头可授性,花粉/胚珠比(P/O),杂交指数(OCI)进行了检测,并进行了人工授粉实验。结果表明:(1)翠雀的花序为总状花序,具3~15朵花,花梗长1.5~3.8cm,单花期10d,花序花期15~18d。(2)翠雀花粉活力日变化呈先升高后降低的趋势;柱头在开花当天至开花后4d无可授性,在开花后的5d、6d有可授性。(3)翠雀的花粉胚珠比(P/O)为2 159.14,杂交指数OCI为5;人工授粉实验表明,翠雀在自然条件下能够结实,结实率为100%;存在自主自花授粉现象,部分自交亲和,结实受传粉者限制;不存在无融合生殖现象。(4)翠雀主要的访花昆虫有四条无垫蜂和短腹管蚜蝇。研究认为,翠雀的繁育系统为兼性异交或专性异交,部分自交亲和,需要传粉者。     

刺山柑雄全同株性系统的适应意义

对分布于新疆北部干旱荒漠区的刺山柑Capparisspinosa性系统进行了研究.结果表明:(1)该物种具有典型的雄全同株性系统.雄花和两性花的雄蕊均正常发育,且有长短之分;两性花的雌蕊发育正常,而雄花的雌蕊败育,只行使雄性功能.(2)居群间每天开放的雄花和两性花比率、两性花的长短雄蕊数及雄花短雄蕊的花丝与花药长等存在极显著差异(P<0.01),但花器官生物量没有显著差异(P>0.05).(3)单花花期为15-16 h,每天18:00时左右开始开放,有强烈的气味和花蜜产生.植株每天产生的雄花和两性花数是随机的,可形成短时的雄花与两性花异株,但居群内雄花数比两性花数多.(4)三个居群两性花的P/O值分别为1.57×104、1.65×104和1.71×104.居群内雄花和两性花的花粉数及长、短雄蕊花药的花粉数均无显著差异(P>0.05),居群间雄花和两性花的花粉数、两性花的胚珠数及P/O值差异也不显著(P>0.05).(5)各居群雄花和两性花长短雄蕊的花粉活力动态变化曲线相似,花粉寿命为18-20 h,两性花雌蕊柱头的可授期为16-18h.(6)访花昆虫为膜翅目Hymenoptera和鳞翅目Lepidoptera昆虫,3个居群共有7种访花昆虫,其活动受天气影响很大.(7)两性花不存在无融合生殖现象,授自花花粉、同株异花花粉和异株异花花粉后均可结实,属于混合交配系统.刺山柑的雄全同株性系统可能是在长期与荒漠环境相适应的过程中由遗传和环境共同作用的结果.雄花的出现不仅增加了花粉数和P/O值,提高了植株的雄性适合度,同时增加了花的数量、对传粉昆虫的吸引力以及两性花柱头接受异花花粉的机会,提高了异交率和雌性适合度,保障其在荒漠极端环境中繁殖成功.     

青藏高原特有植物祁连獐牙菜（龙胆科）的花综合征与虫媒传粉

雌雄异熟和异型花在一些被子植物中的共同存在被认为是一种有效避免两性花植物自交并促进异交的机制。本文报道青藏高原特有植物祁连獐牙菜Swertia przewalskii Pissjauk.的花同时具有雌雄异熟和异型花这两种特征。在此基础上,我们详细观察了该植物访花昆虫的访花行为,主要探讨这种促进异花传粉的机制是否能完全避免株内异花传粉的自交。繁育系统试验证明祁连獐牙菜单花套袋不结实,说明雌雄异熟和异型花的花特征完全避免了单花的自花传粉,自然结实必须依赖昆虫传粉才能完成。然而,该植物却是高度自交亲和的,因为人     

珍稀濒危植物猬实的开花特性与传粉生物学研究

对迁地保护的珍稀濒危植物猬实的开花动态、花粉活力、柱头可授性、访花昆虫种类、访花频率、花粉与胚珠比(P/O)、异交指数(OCI)进行观测.结果表明：猬实的单花期约为7 d,种群花期约为14 d,花冠展开后2~3 h,花粉活力达到最高,约为90％,3 d后花粉基本不具活力.在整个单花期,柱头一直具可授性,花冠展开后第2天可授性最强.花粉与胚珠比(P/O)为(398．1±63．7),异交指数(OCI)为3,表明猬实是以异交为主,自交亲和的繁育系统,有时需要传粉者完成传粉过程.猬实的访花昆虫有10余种,以膜翅目蜜蜂科和隧蜂科昆虫为主,另外还有少量双翅目食蚜蝇科昆虫,偶见鳞翅目的柑橘凤蝶和长喙天蛾,其中以蜜蜂科蜜蜂属昆虫传粉效率最高.迁地保护的猬实未见结籽,其原因还有待于进一步研究.     

野牡丹异型雄蕊的功能分化

野牡丹科植物具有形态、大小和颜色显著不同的两种异型雄蕊。关于其异型雄蕊是否具有功能分化还一直存在争论。本文以野牡丹科植物野牡丹(Melastoma malabathricum)为实验材料, 比较了两种异型雄蕊在传粉过程中的功能作用。结果表明, 两种异型雄蕊在形态、花粉量和人工控制实验条件下的结籽数, 以及主要传粉昆虫木蜂(Xylocopa sp.)访花时的行为等方面都有显著差异, 说明两种雄蕊在传粉过程中存在一定的功能分化: 外轮紫色雄蕊中的花粉为后代提供雄配子, 而内轮黄色雄蕊中的花粉则为传粉昆虫提供食物。但两种雄蕊在花粉活性、花粉组织化学成分和结实率方面差异均不显著, 表明两者在生理上并没有分化。实验结果还表明, 除花前套袋不结实外, 自交、异交和自然对照都具有较高的结实率, 说明野牡丹不存在无融合生殖和主动自交及自交不亲和现象, 为兼性异交。     

Andromonoecy in an Old World Papilionoid legume,Erophaca baetica

Andromonoecy (i.e. the occurrence on individual plants of hermaphroditic and male flowers) is a rare sexual system among the angiosperms, regarded by some authors as a transitional stage from hermaphroditism to monoecy. Having discovered the occurrence of andromonoecy in Erophaca baetica (a Mediterranean shrubby legume with two subspecies), a novelty for Old World papilionoid legumes, we investigated the morpho‐functional correlates and the geographical distribution of this phenomenon in the species. The relative frequencies of hermaphrodite and male flowers were determined in two field and 111 herbarium populations. Biomass allocation within flowers, pollen production and viability, pollen tube growth, nectar production and the temporal pattern of male flower production were also studied in two nearby southern Spanish populations. Virtually all of the studied populations were andromonoecious. Male flowers tended to appear at apical positions within the inflorescence, and became more abundant by the end of the flowering season. Male flowers were externally similar to hermaphroditic flowers (although with less biomass and smaller parts) and released equivalent amounts of pollen and nectar; however, their pollen germinated significantly better. Erophaca is the first example of an andromonecious Papilionoid in the Old World. Since the main difference among floral morphs in this species is functional (i.e. pollen germination rate) rather than morphological, andromonoecy is not readily noticeable, and very careful inspection may be required to reveal it. The potential effect of andromonoecy in enhancing outcrossing rate in this species is discussed.     

Floral biology and pollination mechanisms of four Mexico-endemic Fuchsia species with contrasting reproductive systems

Aims The evolution of the outstanding variation of reproductive systems in angiosperms has been considered an important driver of lineage diversification. Closely related hermaphroditic and dioecious species with biotic pollination provide the opportunity to study and compare traits related to pollinator attraction and their consequences on reproductive components. A higher predictability of pollination syndromes is expected in dioecious species, which are dependent on pollinators, than in self-compatible hermaphroditic taxa. Dioecious species may suffer pollen limitation depending on the quality of floral rewards and the kind and abundance of pollinators, whereas no pollen limitation is expected in hermaphroditic species with autonomous self-pollination. Additionally, in the absence of pollen limitation, more or better seeds are expected in dioecious species, according to the sexual specialization hypothesis.Methods In natural populations of the hermaphroditic Fuchsia fulgens and Fuchsia arborescens and dioecious Fuchsia parviflora and Fuchsia obconica, all endemic to Mexico, we first described flower phenology, flower production and longevity and nectar volume and concentration. Then, we evaluated the correspondence between floral visitors and pollination syndromes. In hermaphrodite plants, we determined the level of herkogamy and the potential for autonomous self-pollination. Finally, we evaluated the effect of pollen limitation on fruit set and seed number and assessed seed germination for all species.Important findings In contrast to our prediction, dioecious species did not show a higher correspondence between pollination syndromes and floral visitors than did hermaphrodites; however, male flowers exhibited a higher correspondence than female flowers. No pollen limitation was detected in dioecious species, for which visitation rate did not differ between male and female flowers. The hermaphroditic F. fulgens showed pollen limitation for seed number, despite the presence of autonomous selfing. Fruit set from autonomous pollination was higher in F. arborescens, which showed a lower level of herkogamy compared with F. fulgens. Finally, dioecious species produced fewer but heavier seeds compared with hermaphrodite species. Although Fuchsia is classified as an outcrossing genus, both hermaphroditic species showed autonomous self-pollination. The heavier but lower number of seeds per fruit in dioecious species may be related to the more efficient resource allocation expected from sexual specialization. This could play an important role in the evolution of dioecy; however, a comparative phylogenetic approach is required to confirm this hypothesis.     

冠果草的性表达状态及其进化含义

通常认为雄花两性花同株是植物性表达两性花进化到雌雄异株的过渡类型之一。慈姑属仅冠果草（Ｓａｇｉｔｔａｒｉａ ｇｕｙａｎｅｎｓｉｓ Ｈ．Ｂ．Ｋ．ｓｕｂｓｐ．ｌａｐｐｕｌａ（Ｄ．Ｄｏｎ）Ｂｏｊｉｎ）的性表达为典型的雄花两性花同株,其他种为雌雄同株。对冠果草野外和实验居群的性配置进行了定量观察。东乡和武夷山居群雄花的比例分别为２．４８％０．９６％。实验表明,栽培条件影响冠果草的性表达：在营养较好的条件下     

SEXUAL ALLOCATION AND ANDROMONOECY: RESOURCE INVESTMENT IN MALE AND HERMAPHRODITE FLOWERS OF SOLANUM CAROLINENSE (SOLANACEAE)

Male and hermaphrodite flowers from the andromonoecious perennial Solanum carolinense were compared. Hermaphrodite flowers were 11–16% heavier than males, and this difference was consistent for two clones studied. Hermaphrodite flowers contained a greater amount of nitrogen than did male flowers, but males had higher percent nitrogen. Male and hermaphrodite flowers had equal investment in male structures: stamen mass, number of pollen grains, mass of pollen grains ejected by vibration, and pollen size were all similar for the two flower types. In contrast, male flowers had reduced masses of disfunctional female structures: ovaries of males were ⅓ as massive, and styles 1/7; as massive, as those of hermaphrodites. These results are consistent with the hypothesis that male flowers (which occur where fruit set is unlikely) increase male fitness through pollen donation, while saving resources that would otherwise be invested in hermaphrodite flowers that abscise.     

Sex Expression and the Evolutionary Advantages of Male Flowers in an Andromonoecious Species, Sagittaria guyanensis subsp. lappula (Alismataceae)

Sex expression in Sagittaria guyanensis H.B.K. subsp. lappula (D. Don) Bojin is typically andromonoecious while the other species in the same genus are basically monoecious. The evolutionary advantages of male flowers and hermaphrodite flowers in S. guyanensis subsp. lappula were assessed by measuring sex allocation and pollen movements in two wild populations of the species. Two cultivated populations served as controls. The percentage of male flowers was very low in the two wild populations in Dongxiang, Jiangxi Province and Wuyishan, Fujian Province, viz., 2.48% and 0.96% respectively. In the two cultivated populations, male flower percentage significantly increased when the soil was of higher nutrient content. This indicates that the allocation to male versus female reproduction might change in response to environmental factors. Pollen production per male flower was 4.1 times higher than that of a hermaphrodite flower. The floral shape and size of male and hermaphrodite flowers were similar. No difference was observed between these dimorphological flowers in pollen germination rate in vitro and in the speed of pollen tube growth in vivo. Anthesis was only 4-5 h. Male flowers usually opened 0.5 h earlier than hermaphrodite flowers. An unexpected finding was that no pollen from the male flowers was found on the stigmas of the hermaphrodite flowers, in spite of the occasional visits by insects to both types of flowers in both wild and control populations. A consistent pattern of fruit development was found to exist in open pollinated flowers as well as in flowers that had been bagged. The sex ratios did not have significant influence on fruit set. Approximately 25% of the pistils in a gynoecium failed to develop into fruits because no pollen was deposited on them, indicating that the fruit set of this andromonoecious plant is mainly affected by pollen limitation rather than resource limitation. Reproduction in S. guyanensis subsp. lappula in the habitats was dependent on self-pollination in hermaphrodite flowers. The male flowers in this species might be a potential source of additional pollination and may facilitate cross-pollination. The fact that the flowers of monoecious species in Sagittaria pollinated by a wide diversity of insect visitors may contribute enormously to the diversification of sex expression in this genus.     

PHENOTYPIC GENDER IN SEX CHANGING DWARF GINSENG,PANAX TRIFOLIUM (ARALIACEAE)

Dwarf ginseng (Panax trifolium L., Araliaceae) is a diphasic (“sex changing”) species in which one phase has staminate flowers and the other has hermaphroditic flowers. In order to determine the relative allocations of the hermaphroditic gender phase to male and female functions,variation in population gender phase ratios, pollen production and viability, and ovule and seed production were documented. Gender phase ratios are highly male-biased. Dwarf ginseng is self-compatible, and both gender phases have viable pollen capable of effecting fertilization. Males produce more flowers and more viable pollen per anther than hermaphrodites. The phenotypic gender of hermaphrodites is extremely female-biased; it is likely that hermaphrodites function essentially as females. Sexual selection may have a role in the evolution and maintenance of differences between the gender phases in allocation to male function.     

Pollen viability and limitation of seed production in a population of the circumpolar cushion plant, Silene acaulis (Caryophyllaceae)

Pollen viability among genders and limitation of female seed production in a natural trioecious population of the circumpolar cushion plant Silene acaulis was examined. Pollen viability was estimated by an in vitro pollen germination experiment. Both male and hermaphrodite flowers displayed large variation in pollen viability (0–53% in hermaphrodite and 0–54% in male flowers). There was a significant difference between genders in pollen viability: male plants had on average higher pollen viability than hermaphrodite plants. Resource and pollen limitation of seed production was studied by an experiment consisting of three treatments; (I) hand-pollination and removal of all other flowers on the cushion, (II) hand-pollination without removal of other flowers, and (III) open pollination without removal of flowers. Hand-pollination increased seed production, whereas removal of flowers had no effect on seed production. Abortion of pollinated ovules during seed development and seed mass did not differ among treatments. To control for effect of fruit number on seed production, data from naturally pollinated individuals was used. There was a positive correlation between both total number of seeds and fruit number, mean seed number per fruit and fruit number, respectively. These results indicate that seed production of 5. acaulis is mainly limited by pollen availability whereas resource competition between fruits is not important as a limiting factor. The possible role of male quality differences between genders and pollen limitation of seed production for maintenance of trioecious reproductive systems is discussed.     

Size-dependent gender modification in Lilium apertum (Liliaceae): does this species exhibit gender diphasy?

Background and Aims

Variation in the relative female and male reproductive success of flowering plants is widespread, despite the fundamental hermaphroditic condition of the majority of species. In many hermaphroditic populations, environmental conditions and their influence on development and size can influence the gender expression of individuals through the formation of hermaphroditic and unisexual flowers. This study investigates the hypothesis that the bulbous, animal-pollinated, perennial Lilium apertum (Liliaceae) exhibits a form of size-dependent gender modification known as gender diphasy, in which the sexual expression of individuals depends on their size, with plants often changing sex between seasons.

Methods

Variation in floral traits was examined in relation to their size using marked individuals in natural populations, and also under glasshouse conditions. Measurements were taken of the height, flower number, floral sex expression, flower size, flower biomass and pollen production of individuals over consecutive years between 2009 and 2012 in seven populations in south-west China.

Key Results

Flowers of L. apertum are either perfect (hermaphroditic) or staminate (male) and, in any given season, plants exhibit one of three sex phenotypes: only hermaphrodite flowers, a mixture of hermaphroditic and male flowers, or only male flowers. Transitions between each of these sex phenotypes were observed over consecutive years and were commonly size-dependent, particularly transitions from small plants bearing only male flowers to those that were taller with hermaphroditic flowers. Hermaphroditic flowers were significantly larger, heavier and produced more pollen than male flowers.

Conclusions

The results for L. apertum are consistent with the ‘size advantage hypothesis’ developed for animal species with sex change. The theory predicts that when individuals are small they should exhibit the sex for which the costs of reproduction are less, and this usually involves the male phase. L. apertum provides an example of gender diphasy, a rare sexual system in flowering plants.     

不同海拔藜芦种群繁殖特征的初步研究

 在北京东灵山地区,沿一定海拔梯度,对藜芦（Veratrum nigrum）这一雄花两性花同株的多年生草本植物的繁殖特征进行了初步研究。结果表明：藜芦开花植株的生物量显著大于未开花植株的生物量。藜芦开花植株大小存在最小临界值,总种群的最小临界值为2.61 g,不同种群的繁殖临界值存在一定的差异。根据花粉/胚珠比（Pollen / ovule ratio,P/O比）推测,藜芦应属于以异交为主的混合交配系统。雄花的大小与生物量都比两性花小,而且其开放时间也晚于两性花,但二者的花粉生产量却没有显著差异。这种现象与人们针对雄花两性花同株植物的雄花功能所提出的最优资源分配假说（Optimal resource allocation hypothesis）是一致的。     

Pollen Production in the Androdioecious Datisca glomerata (Datiscaceae): Implications for Breeding System Equilibrium

Abstract Androdioecy (the presence of males and hermaphrodites in a breeding population) is a rare reproductive system in plants, with Datisca glomerata (Datiscaceae) representing the only well-documented example. Recent reports of high outcrossing rates, inbreeding depression, and high male pollen production satisfy theoretical predictions for the continued maintenance of androdioecy in populations of this species. However, in prior studies pollen production was measured indirectly in terms of numbers of anthers per flower—based on the assumption that male and hermaphroditic plant have equal numbers of flowers and that anthers from the two sexual morphs produce equivalent amounts of pollen. Herein, we demonstrate that male and hermaphrodite plants do not differ significantly in terms of flower number, but that pollen production in anthers from hermaphroditic plants is 12.6% higher than in anthers from male plants, thus refining the estimate of relative pollen fecundity of male versus hermaphrodite plants. The differential lowers the frequency of males predicted by theory, but is still consistent with the maintenance of androdioecy in this species.