FUNCTIONAL DIOECY AND ANDROMONOECY IN SOLANUM

Field and laboratory studies of 19 diclinous species endemic to Australia help to clarify the nature and evolution of andromonoecy, androdioecy, and dioecy in the genus Solanum. Ten species are andromonoecious; typically these species bear inflorescences with a single, large basal hermaphroditic flower and 12–60 distal, smaller staminate flowers. We suggest that the andromonoecious condition was derived from hermaphroditic-flowered ancestors in part by hemisterilization of flowers but largely by addition of staminate flowers. The resultant larger inflorescences are hypothesized to serve both to attract and to entrain pollinators, yielding more or higher-quality seed set in hermaphroditic flowers and/or greater dispersion of pollen from staminate flowers. We suggest that andromonoecy may also serve to reduce selling. Nine other species are morphologically androdioecious but functionally dioecious. In these species, staminate flowers, like those of the andromonoecious species, bear anthers with copious tricolporate pollen and a highly reduced gynoecium. The morphologically hermaphroditic flowers are functionally pistillate and borne singly in inflorescences, and they bear anthers with inaperturate pollen. The inaperturate pollen, although viable, never germinates and is hypothesized to be retained in pistillate flowers as a reward to pollinators in the nectarless Solanum flowers. All other species of Solanum studied with pollen dimorphism in which one pollen morph is inaperturate are also best treated as functionally dioecious. We conclude that there is no evidence for androdioecy in Solanum. A review of other families suggests that there is little support for this unusual breeding system in any other angiosperm group either. Preliminary analyses suggest that andromonoecy and dioecy are polyphyletic in Solanum. Furthermore, dioecy is as likely to have arisen from hermaphroditic as from andromonoecious ancestors.     

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。     

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.     

Systematic significance of pollen nucleus number in the genus Saxifraga (Saxifragaceae)

Pollen cell number was studied in 124 species of Saxifraga and six species of Micranthes. Binucleate pollen is present in all the six species of Micranthes studied and in 96 species of Saxifraga. Trinucleate pollen occurs in 28 species of Saxifraga and is confined to section Ciliatae subsection Hirculoideae, where it is a putative synapomorphy, and to two species of subsection Serpyllifoliae. Trinucleate pollen does not characterise the whole of section Ciliatae as has been thought. The state has probably evolved at least twice, and possibly three times, in the genus. One new nomenclatural combination is made in Micranthes.     

Pollen morphology of families Quillajaceae and Surianaceae (Fabales)

The Fabales clade comprises four families: Leguminosae, Polygalaceae, Quillajaceae and Surianaceae. This study presents new information on the pollen morphology of Quillaja, the only genus of Quillajaceae, and Recchia, Guilfoylia, Cadellia, Suriana and Stylobasium, the five genera that comprise Surianaceae. The pollen of 9 of the 11 species currently recognised within the two families was examined using light microscopy (LM), scanning electron microscopy (SEM) and, selectively, with transmission electron microscopy (TEM). Pollen of all taxa is isopolar with tri-zonocolporate apertures, lalongate endoapertures with fastigia adjacent to the endoaperture, and long ectoapertures that are nearly equal to the polar length. Apocolpia are correspondingly small. Quillaja pollen is subprolate to prolate, and striate with a granular aperture surface membrane. Ectexine protrudes over the endoapertures. In thin section the foot layer is thicker in mesocolpial areas and thin to discontinuous around the apertures, where the endexine is thicker. Cadellia pollen is prolate spheroidal, and striate with a granular aperture surface membrane. Exine protrudes over the endoapertures. In thin section the endexine is thicker and lamellate around the endoaperture area, and the foot layer is thicker in mesocolpial regions. Guilfoylia pollen is oblate and gemmate-verrucate, with a granular aperture surface membrane. Columellae are short. Recchia pollen is suboblate to oblate spheroidal, and microreticulate-perforate with a granular aperture surface membrane. Exine protrudes over the endoapertures. The foot layer is thin to discontinuous around aperture margins and thick in mesocolpial regions. Stylobasium pollen is suboblate, and finely rugulate-perforate with a granular aperture surface membrane. Columellae are short, the foot layer is thin or absent. Suriana pollen is suboblate, and finely rugulate-perforate with a granular aperture surface membrane. Pollen of Cadellia and Recchia, and Stylobasium and Suriana are morphologically similar. Verrucate surface ornamentation is only present in Guilfoylia. Quillaja, Cadellia and Recchia share the character of protruding exine over the endoaperture area. Striate ornamentation occurs in Quillaja and Cadellia. The pollen morphology of Quillajaceae has more in common with that of Leguminosae and Surianaceae, and with Cadellia in particular, than with Polygalaceae.     

Functional Monoecy Due to Delayed Anther Dehiscence: A Novel Mechanism in Pseuduvaria mulgraveana (Annonaceae)

Unlike most genera in the early-divergent angiosperm family Annonaceae, Pseuduvaria exhibits a diversity of floral sex expression. Most species are structurally andromonoecious (or possibly androdioecious), although the hermaphroditic flowers have been inferred to be functionally pistillate, with sterile staminodes. Pseuduvaria presents an ideal model for investigating the evolution of floral sex in early-divergent angiosperms, although detailed empirical studies are currently lacking. The phenology and pollination ecology of the Australian endemic species Pseuduvaria mulgraveana are studied in detail, including evaluations of floral scent chemistry, pollen viability, and floral visitors. Results showed that the flowers are pollinated by small diurnal nitidulid beetles and are protogynous. Pollen from both hermaphroditic and staminate flowers are shown to be equally viable. The structurally hermaphroditic flowers are nevertheless functionally pistillate as anther dehiscence is delayed until after petal abscission and hence after the departure of pollinators. This mechanism to achieve functional unisexuality of flowers has not previously been reported in angiosperms. It is known that protogyny is widespread amongst early-divergent angiosperms, including the Annonaceae, and is effective in preventing autogamy. Delayed anther dehiscence represents a further elaboration of this, and is effective in preventing geitonogamy since very few sexually mature flowers occur simultaneously in an individual. We highlight the necessity for field-based empirical interpretations of functional floral sex expression prior to evaluations of evolutionary processes.     

Morphology and variability of pollen of the genus <Emphasis Type="Italic">Spiraea</Emphasis> L. (Rosaceae) in Siberia and the Far East

Pollen of 12 species of the genus Spirea L. growing in different regions of Siberia and the Far East was investigated. The ultrasculpture of pollen grains is described. On the basis of aperture morphology and exine sculpture, pollen grains are typified, with key indices given for their identification. The species of the Spiraria section are distinguished by large grains. Pollen grains of S. media and S. alpina considerably vary in size. The level of natural polymorphism of palynological indices has been established.     

New insights into the pollen morphology of the genus Mutisia (Asteraceae, Mutisieae)

The pollen morphology from 67 collections representing 52 species of Mutisia was analyzed by means of light, scanning electron and transmission electron microscopy. Pollen of Mutisia is comparable to that of the more widespread type found in Mutisiinae. The pollen grains of the genus are characterized by the spheroidal-subprolate to prolate shape, large size, tricolporate aperture, with mesoaperture, exine Mutisia type, with sculpturing microechinate to rugulate or microechinate-rugulate. This basic pollen type is very variable in size, endoaperture shape, and exine thickness of the pollen grains. A new type of arrangement of columellae was observed in pollen of Mutisia section Mutisia (e.g., M. stuebelli, M. microphylla, and M. sodiroi). It is suggested that it might have harmomegathic significance. The pollen morphology and polymorphism are discussed with regard to functional mechanisms and possible evolutionary significance.     

Pollen heteromorphism is pervasive in <Emphasis Type="Italic">Thalictrum</Emphasis> (Ranunculaceae)

Among the angiosperms, features of pollen morphology such as grain size, aperture number and surface ornamentation display striking variation. It is less well appreciated that pollen morphology may vary within and among populations of the same species as well as within individual plants. In some species, individual plants produce multiple types of fertile pollen grains (called pollen heteromorphism). Aspects of pollen morphology, such as aperture number, are likely to affect fertilization success with different morphologies favored in different local competitive and ecological environments. This study surveys variation in pollen grain morphology among species throughout the genus Thalictrum. Pollen from individuals of 36 species was rehydrated from herbarium specimens, and light microscopy was used to quantify pollen grain aperture number and size. I find that pollen aperture-number heteromorphism is present within all Thalictrum species studied, and distributions of aperture-number morphs vary both within and among species. This study provides an example of significant pollen heteromorphism within a genus that also varies widely for pollination mode and sexual system.     

Pollen morphology of the Pavetteae (Rubiaceae,Ixoroideae) and its taxonomic significance

Pollen grains of the tribe Pavetteae (Rubiaceae, subfamily Ixoroideae) are examined using LM and SEM. Grains are 3‐ or 4‐colporate and (semi‐) tectate (in one Versteegia species atectate). Sexine patterns vary between perforate, microreticulate, reticulate, rugulate and striato‐reticulate. Supratectal elements are sometimes present. The variation in pollen morphology in the Pavetteae allows to recognize seven pollen types, the distribution of which is useful to evaluate generic delimitations and relationships within the tribe. Pollen characters corroborate the close relationships between the genera Coleactina, Dictyandra and Leptactina and between Homollea, Homolliella and Paracephaelis. All the genera of the tribe proved to be stenopalynous (the species examined possess the same pollen type), except Pavetta, Rutidea, Versteegia and Tarenna which are eurypalynous. In the huge genus Pavetta the existing infrageneric classification is supported pollen morphologically. Pollen morphology further indicates that the genus Tarenna is badly delimited and strongly in need of a revision. The small genus Versteegia is in need of further taxonomic and palynological study to understand the pollen morphological variation encountered here. At a higher rank, pollen morphology also does not contradict the recent division of the Pavetteae in the Ixoreae (a stenopalynous tribe with presumably primitive pollen) and the Pavetteae sensu stricto (eurypalynous).     

POLLEN MORPHOLOGY AND SYSTEMATICS OF SIPHONOGLOSSA SENSU LATO (ACANTHACEAE)

Pollen morphology of 15 species of Siphonoglossa and of two closely related groups was investigated. Two tribal-specific pollen types are found within Siphonoglossa sensu lato suggesting that the genus is artificial, composed of taxa belonging in several genera among two tribes (subtribes sensu Bremekamp) of Acanthaceae. Five taxa currently included in an informal subgeneric category of Siphonoglossa have tricolporate, prolate pollen (termed Type I) that is characteristic of Odontonemeae (= Odontoneminae, Justicieae). Pollen of the remaining taxa, belonging in two formal sections of the genus, are mostly 2-porate, bilateral (Type II) with a sexine sculpturing characteristic of Justicieae (= Justiciinae). Pollen of section Siphonoglossa is rather uniform, 2-porate, bilateral with lolongate pores, and seem to delimit a natural group. Taxa of section Pentaloba have a more heterogeneous pollen morphology, mostly 2-porate, bilateral with lalongate pores. Controversial aspects of the interpretation of pollen morphology in Justiciinae are presented and their relevance to this study are examined. Hypothetical trends in the evolution of pollen of Justiciinae are discussed and the application of pollen morphology to taxonomy of the genus is presented, including a recommendation for narrowing the generic concept of Siphonoglossa to the taxa of the type section.     

COMPARATIVE REPRODUCTIVE BIOLOGY OF ACER PENSYLVANICUM AND A. SPICATUM (ACERACEAE)

Pollination mechanisms and floral features affecting reproduction were examined in two closely-related species, Acer pensylvanicum and A. spicatum. Intra-specific crossing experiments, studies of pollen stainability and ovule number, and observations of the flowers in the field revealed that neither species bears functionally hermaphroditic flowers. Three flower-types were found in each species: staminate, pistillate, and a third type that is morphologically hermaphroditic. The latter form sheds viable pollen, but the pistil does not contain ovules. Pollen-ovule ratios of both species are in the range of expectation for a facultatively xenogamous reproductive system. Outcrossing is accomplished by entomophily in both species. A variety of insects from several orders visit the flowers; however bees in the genus Andrena (Hymenoptera: Andrenidae), especially A. milwaukeensis, are considered the most important pollinators of both maple species because of their fidelity to Acer, their intrafloral behavior, and their ability to carry pollen. However, flies are the most abundant visitors to A. pensylvanicum. Apomixis is an important means of reproduction in A. spicatum. Although floral phenology of these maple species overlaps within a geographical region they were never found blooming simultaneously when growing together.     

Comparative pollen morphology of annual Trigonella L. (Fabaceae) in Turkey

Pollen morphology of 38 Trigonella species was investigated with light microscopies and scanning electron microscopies. Pollen slides were prepared using Wodehouse technique. The pollen grains of Trigonella members are radially symmetrical and isopolar, their outlines are oblong in equatorial view and circular in polar view. Amb is semicircular. The pollen grains are prolate-spheroidal, oblate-spheroidal, suboblate, spheroidal, and prolate with the polar axes 13.5–49.9 μm and the equatorial axes 10.4–43.7 μm. The smallest dimensions are observed in T. spinosa and T. brachycarpa, and the largest in T. carica and T. rhytidocarpa. The pollen grains of Trigonella taxa are usually tricolporate or rarely tricolpate, pantocolporate, tetracolporate, and tricolpodiporate. Various ornamentation types were observed: Microreticulate (section Samaroideae, Pectinatae, Falcatulae, Cylindricae, Bucerates, Reflexae, Isthmocarpae, Capitatae and Foenum-graecum), rugulate-scabrate (section Lunatae, Uncinatae), perforate (T. halophila, T. rigida and T. carica from section Bucerates) and retipilatae (section Biebersteinianae) at equator region and psilate-perforate (section Samaroideae, Pectinatae, Bucerates, Reflexae, Isthmocarpae, Foenum-graecum, T. coelesyriaca, T. cilicica), rugulate-scabrate (section Lunatae, Uncinatae), microreticulate (Falcatulae, Cylindricae and Foenum-graecum) and retipilatae (section Biebersteinianae) at polar region. Ornamentations, pollen shape and the aperture type have been observed as important morphological characters.     

Shared traits make flies and bees effective pollinators of oilseed rape (Brassica napus L.)

One of the major aims of pollination ecology has been to understand the role and relative importance of different pollinator species in both natural and agricultural systems. This study explores how the quantity of pollen delivered in a single visit to a stigma (SVD) differs across insect species from a wide range of taxa (Orders: Hymenoptera, Diptera and Coleoptera) and assesses the morphological and behavioural traits that lead to differences in pollen delivery. We used oilseed rape (OSR) (Brassica napus L.), an economically important crop with open self-fertile flowers, as a test system. Behavioural traits and SVD were measured in the field for individuals of all frequent flower-visiting species of OSR. Individuals were collected and morphological traits were measured in the laboratory. The quantities of free pollen on different body parts of flower visitors, or ‘pollen load’, were also measured. Behavioural and morphological traits were then assessed as predictors of SVD. The individuals that delivered most pollen were (in order of greatest median number of pollen grains delivered): bumblebees (Bombus spp.), mining bees (Andrenidae) and honeybees (Apis mellifera). However, all but one family of flies tested delivered a significantly greater median number of pollen grains than the control, and most delivered more than sweat bees (Halictidae). Behavioural and morphological traits were important in determining pollen delivery, with greater body length, hairiness and visit duration all resulting in greater SVD. Pollen load was measured for a proportion of individuals and we found that for momentary flower visits, a greater quantity of pollen on the head of the flower visitor was linked to a greater SVD. This study demonstrates that, while bees are important pollinators in this system, many fly taxa are also effective at delivering pollen and this is linked to morphological and behavioural traits found in both groups.     

Relationships of pollen size,pistil length and pollen tube growth rates in Rhododendron and their influence on hybridization

Summary Pollen size and pistil length data have been collected for 93 species of Rhododendron (Ericaceae) belonging to a number of different subgeneric taxa. For a sample of eight species in section Vireya, pollen tube growth in the style after selfor interspecific pollination has been quantified. Pollen volume and the time taken for pollen tubes to reach the ovary were both related to pistil length. Pollen-tube growth rates were generally greater for species with longer pistils and larger pollen. Increasing temperature increased the rate of pollen-tube growth. There was no detectable effect of pollen tube density on tube growth rate in the style. After interspecific pollinations tube growth rates in foreign styles could be faster or slower than in self styles. A semisterile individual with two viable pollen grains per tetrad and a plant grafted as scion to a longer-styled stock both showed more rapid pollen-tube growth than expected on the basis of pistil size. Data collected for 26 species in section Vireya showed that where extreme disparity of pollen/pistil size causes failure of interspecific crosses, one or more bridging species with intermediate pollen/pistil size can generally be selected.     

Influence of temperature on the in vitro pollen germination and pollen tube growth of various native Iranian almonds (Prunus L. spp.) species

Pollen germination and pollen tube growth was quantified among various native Iranian wild almonds (P. dulcis (Mill.) D. A. Webb, P. eleaegnifolia Mill., P. orientalis Mill., P. lycioides Spach, P. reuteri Bioss. et Bushe, P. arabica Olivier, P. glauca Browick and P. scoparia Spach in order to identify differences in the tolerance of pollen to temperature variations. Pollen germination and pollen tube growth were observed after incubation in darkness in a germination medium for 24?h at 10?C50°C at 5°C intervals. Maximum pollen germination of the wild almond species and specify that 60% was obtained for P. orientalis pollen and 98% for P. scoparia. Pollen tube length ranged from 860???m was obtained in P. lycioides and 1490???m in P. scoparia. A modified bilinear model best described the response to temperature of pollen germination and pollen tube length. Almond species variation was found for cardinal temperatures (T _min, T _opt and T _max) of pollen germination percentage and pollen tube growth. Mean cardinal temperatures averaged over eight almond species were 14.7, 24.2, and 43.7°C for maximum percentage pollen germination and 14.48, 25.3, and 44.4 °C for maximum pollen tube length. The principal component analysis (PCA) identified maximum percentage pollen germination and pollen tube length of the species, and T _max for the two processes as the most important pollen parameters in describing a species tolerance to high temperature. PCA also classified Prunus L. spp. into four groups according to the tolerance of pollen to temperature variations. The T _min and T _opt for pollen germination and tube growth, rate of pollen tube growth were less predictive in discriminating species for high temperature tolerance.     

Effect of calcium and sucrose concentration on pollen germination invitro of sixRosa species

The germination of the pollen from11 individuals of six wildRosa species was studied. The presence of calcium resulted in increased pollen germination, longer pollen tubes, and in a decrease of the requirement of sucrose concentration in cultivation media. Pollen germination in the medium with Ca reached the values of pollen viability estimated by tetrazolium test in all cases except with roses with balanced heterogamy, of the sectionCaninae. The stimulating effect of calcium was generally most pronounced in the pollen from roses of hybrid nature, such asR. jundzillii, R. canina, and especially in the case of the calciphilous speciesR. eglanteria.     

Pollen morphology of Astragalus section Hymenostegis (Fabaceae) and evaluation of its systematic implications

Astragalus is with nearly 3000 described species the largest genus of flowering plants. So far analyses of pollen characters have only been conducted for a few species of the groups within the genus. Here we analyse pollen grains of 22 species representative for Astragalus section Hymenostegis using scanning electron microscopy. We found the basic shape of the pollen grains to be oblate-spheroidal and apertures to be tricolpate as for other eudicots. The sculpturing pattern of the exine is micro-reticulate. Pollen grains show low morphological variation among different species of this section, but differences occur between sections of the genus. We conclude that the vast morphological differentiation that occurred during the rapid radiation of section Hymenostegis was not accompanied by comparable differentiation in pollen morphology.