Pollen morphology in thePortulacaceae (Centrospermae)

Pollen of 110 species from 18 genera in thePortulacaceae has been examined by light and scanning electron microscopy, and a representative number by transmission electron microscopy. Three basic pollen types were found: 3-colpate with thick tectum and foot layer with prominent unbranched columellae and an extremely thin endexine; pantoporate with thick tectum and foot layer with branched columellae enclosing pores and an endexine that is one to two layers thick; pantocolpate with thin tectum and foot layer with broad, short unbranched columellae and an inconspicuous endexine. All pollen types, however, have a spinulose and tubuliferous/punctate ektexine. Also, all the genera except three,Calandrinia H.B.K.,Montia L. andTalinum Adanson are stenopalynous. There is, however, no absolute correlation between pollen morphology and geographical distribution, although both the major centre of palynological diversity and the majority of all species with tricolpate grains occur in South America.     

Caractères ultrastructuraux du pollen de quelques Podostémales. Affinités avec les Rosidae évoluées

The order Podostemales are two pantropical families of aquatic plants living in running water: Tristichaceae (five genera, ten species) and Podostemaceae (35 genera, 200 species). Pollen of the 26 genera and 62 species studied is characterized by: a granular infratectum in which the granules are sometimes organized as columellae like units, and a lamellar and/or granular endexine in all pollen types, single or in dyads. Most of the apertural characters and the ornamentation of the exine allow us to distinguish the previously established taxonomic groups: Weddellinoideae have tricolporate rugulo-areolate, pollen with a smooth apertural membrane; in Tristichoideae, pollen is periporate and the microspines of the tectum and of the apertural membrane are massive; in Podostemaceae, the tectal spines with their broad bulbous base are formed from numerous masses of sporopollenin, the endexine is microfibrillar at the base, the apertural membrane is constituted of structured ectexinous masses, and the endexine is granular at the aperture. Presence of the tricolporate pollen type associates the Podostemales with the higher eudicotyledons. Most of the pollen characters of Podostemales and their variations are found among advanced Rosidae (Hamamelidales, Polygalales, Santalales, Violales, Euphorbiaceae).     

Wall development and its autofluorescence of sterile and fertileVicia faba L. Pollen

Summary The ultrastructural changes of the pollen wall of three types of fertile and one of sterileVicia pollen were related to the autofluorescence of the pollen wall, measured by a microspectroscopic method. Till the liberation of the microspores from the tetrad, the spectrum of the ectexine shows sometimes two maxima and has a very low intensity. After this period the endexine is formed and its spectrum has one maximum with a high intensity. The differences of the pollen wall between the sterile and fertile pollen exist of the presence of one spectral maximum during the tetrad stage, a thick endexine and the absence of the intine in the sterile pollen. The different types show much differences during the tetrad stage in the callose wall as well as the ectexine. The autofluorescence illustrates the complexity and specificity of the pollen wall development.     

国产蔷薇科李亚科的花粉形态

本文报道了国产李亚科（Prunoideae）10属11种植物花粉形态。花粉近球形至长球形,极轴 20.00－44.12um,赤道轴门17.85-36.95um。极面观三裂圆形,赤道面观椭圆形至圆形。具三孔沟,内孔常为长方形。沟较长,两端较窄。但在 Sinoplagiospermum uniflora中除了3孔沟外,还有周孔沟。花粉外壁明显分化为覆盖层和柱状层,在光学显微镜下表面常模糊,扫描电镜下外壁纹饰均为条纹状。抱粉学结合形态学和细胞学证据说明本亚科为一单系发生的类群。Prinsepia utilis 和Sinoplagiospermum uniflora（Prinsepia uniflora）这两个种在花粉特征和外壁纹饰上差异很大,从而支持将 Prinsepia和 Sinopmplagiospermum分别处理为两个属但不支持将广义的Princepia（含Princepia和Sinoplagiospermum）独立为亚科。此外,由于Exochorda的外壁纹饰同Prunoideae中较原始的类群（如 Laurocerasus）相近,结合细胞学和形态学证据,支持将 Exochord移置至Prunoidea之下。     

五列木科和肋果茶科花粉外壁超微结构及其与山茶科的系统学关系

借助光学显微镜、扫描电镜和透射电镜对五列木科Ｐｅｎｔａｐｈｙｌａｃａｃｅａｅ和肋果茶科Ｓｌａｄｅｎｉａｃｅａｅ花粉进行了观察,并与山茶科Ｔｈｅａｃｅａｅ若干属的花粉进行了详细的比较,同时参考了猕猴桃科Ａｃｔｉｎｉｄｉａｃｅａｅ的花粉特征。研究表明,肋果茶科的花粉与山茶科的花粉具有较多的相似性,如其形状和大小与山茶科中厚皮香亚科的几乎一致;而花粉的表面纹饰（粗糙至模糊的皱波状纹饰）则与山茶属,Ｆｒｅｚｉｅｒａ属和厚皮香亚科部分属的花粉纹饰相类似。从花粉外壁的结构看,肋果茶的花粉与石笔木属和山茶属的更接近,表现为：花粉外壁层次分化明显,复盖层和柱状层均较厚,外壁内层薄。而五列木科花粉除了形状和萌发孔类型与山茶科的相似外,其纹饰特征和外壁结构均与山茶科的差异较大,如五列木科花粉表面近光滑,外壁覆盖层较簿,柱状层很不发达,外壁内层相对较厚等。孢粉学上认为,肋果茶科与山茶科具有更为密切的关系,五列木科则与山茶科较疏远。支持把肋果茶科作为一个属置于山茶科内或作为山茶科的一个亚科     

A light, scanning electron, and transmission electron microscopic study of pollen wall architecture in the subtribe Gnaphaliinae (Gnaphalieae, Asteraceae)

To better understand relationships within the Asteroideae, the pollen morphology and exine structure of 10 genera and 15 species of Gnaphaliinae were investigated using light, scanning electron, and transmission electron microscopy. All taxa have a Gnaphalioid pattern of exine with an evidently rough foot layer. The tectal complex consists of three main layers that differ in morphology and thickness: a tectum, a median columellar layer, and an internal interlaced sub-columellar layer. The apertural system consists of an ectoaperture, a mesoaperture, and an endoaperture, which intersect, respectively, the tectal complex, the foot layer and the upper part of the endexine, and the inner part of the endexine. On the basis of pollen characteristics, especially those of the spines, all the species examined may be gathered into two groups, one including Helichrysum foetidum, H. italicum, Plecostachys serpyllifolia, and Pseudognaphalium luteo-album and the other including the other 11 species of Gnaphaliinae now investigated. In addition, because Gnaphalieae has been proposed as sister group to several tribes and clades of Asteroideae, a comparison was made between exine pattern in the Gnaphalieae and that in its putatively related tribes. On the basis of previous phylogenetic studies and our present pollen data we suggest that either Astereae, Astereae-Anthemideae clade, or Heliantheae s.l.-Anthemideae clade are the best candidates for the sister taxon/sister group of Gnaphalieae.     

Pollen morphology in Campanulaceae IV

The exine structure has been studied in Campanulaceae s. lat. The results are combined with those in previous studies of shape, sculpturing and aperture conditions. Fifteen further species have been studied by SEM. Two main groups of pollen are found: (1) porate pollen with spinules and ridges/protrusions or a low relief reticulum, combined with an ektexine varying from simple to complex and a lamellated endexine; (2) 3–colpate/colporate pollen with a high relief reticulate/striate surface sculpturing, in general a homogeneous ektexine and an endexine lacking lamellae. The first group corresponds to Campanulaceae s. str., the second to Lobeliaceae s. str. Some genera like Cyananthus, Codonopsis and Parishella have unique characters which make them difficult to place in any of the two main groups.     

Crossability Barriers in Neotropical Heliconia

Artificial hybridization among species of neotropical Heliconiawas studied at two sites in Costa Rica, centralAmerica. At LasCruces Tropical Botanical Garden individuals in cultivationwere used as parents in crosses primarily between species withpendent inflorescences that normally are distributed allopatrically.At Finca La Selva normally sympatric species with either pendentor erect inflorescences were crossed in their natural habitats.Observation of pollen tube growth by means of fluorescence microscopyand seed set were used to determine the extent of crossability.Crossability barriers between the majority of species are strongand foreign pollen tubes are inhibited at the stigmatic surface,within the stylar tissue or within the ovary. The site of inhibitionis consistent for each pair of species, and is dependent onthe parentage and the direction of the cross. Although additionalisolating mechanisms, such as pollinator specificity and phenologicalseparation, are present in Heliconia, pre-fertilization crossabilitybarriers act as the ultimate mechanism to prevent hybridization.The type of barrier (stigmatic, stylar or ovarian) that existsbetween two species is not dependent upon the geographical distributionof the parental species or the specific types of pollinatorsthat visit them, but in some cases may indicate taxonomic relationships. Heliconia spp, isolating mechanisms, crossability barriers, progamic phase, hybridization, Costa Rica, hummingbirds, taxonomy, pollinator sharing     

Comparative pollen morphology of Glechoma and Marmoritis(Nepetinae,Lamiaceae)

The pollen morphology of 13 taxa(34 specimens)of the genera Glechoma L.,and Marmoritis Benth.was investigated in detail using light,scanning electron,and transmission electron microscopy.Pollen grains of all studied taxa are small to large in size(P = 32.5-60.4μm,E = 20.2-50.5μm),prolate-spheroidal to prolate in shape and mostly hexacolpate(the amb more or less circular or rarely ellipsoid)with granular membranes.The sexine ornamentation of Glechoma is bireticulate; the muri of the primary reticulum are irregularly circled,and lumen size is short.In contrast,the sexine surfaces of the Marmoritis pollen tend to more elongate or wider at the muri of the primary reticulum than those of the Glechoma.The pollen wall stratification of selected taxa(three from Glechoma and one from Marmoritis)is characterized by unbranched columellae,and continuous or distinctly discontinuous endexine based on transmission electron microscopy observation.The results of Glechoma and Marmoritis reveal rather similar pollen morphological features,however,fine details of sexine ornamentation are characteristic to differentiate the pollen taxa.Although these differences may be useful in establishing the taxonomic boundary between two genera,they are too weak to segregate diagnostic characters.     

The morphology and ultrastructure of Jurassic in situ ginkgoalean pollen

A fragmentary pollen organ with four to six microsporangia is discovered from the Middle Jurassic of the Irkutsk coal basin, Siberia. The in situ pollen grains are boat-shaped, monosulcate, and with a nearly psilate surface. The non-aperture ectexine is composed of a thick solid tectum, a thin infratectum, and a thin foot layer. The infratectum includes one row of small rare alveolae. The supposedly poorly preserved endexine is thin and grainy. The ectexine reduces greatly in the aperture area, where only homogeneous ectexinal patches are present over the endexine. The pollen grains under study resemble in their exine ultrastructure pollen grains of the modern Ginkgo biloba and pollen grains from dispersed seeds of a presumably ginkgoalean affinity from the Middle Jurassic of Uzbekistan. This suggests that the ginkgoalean exine ultrastructure of the modern type existed as early as the Middle Jurassic. The exine ultrastructure under study is also similar, though to lesser degree, to that of dispersed pollen grains of a presumed ginkgoalean affinity from the Cretaceous of the Russian Far East. The diversity of such a long-living group as ginkgoaleans is apparently reflected in the diversity of their exine ultrastructure. To the present knowledge, ginkgoalean pollen grains can be differentiated from similar boat-shaped monosulcate pollens by the following co-occurring characters: a thick homogeneous tectum, a thin infratectum with one row of structural elements, a thin foot layer, and an ectexine that is reduced in the aperture region to patches.     

Comparative exine development from the post-tetrad stage in the early-divergent lineages of Ranunculales: the genera <Emphasis Type="Italic">Euptelea</Emphasis> and <Emphasis Type="Italic">Pteridophyllum</Emphasis>

Studies of pollen wall development produce a great deal of morphological data that supplies useful information regarding taxonomy and systematics. We present the exine development of Euptelea and Pteridophyllum, two taxa whose pollen wall development has never previously been studied using transmission electron microscopy. Both genera are representatives of the two earliest-diverging families of the order Ranunculales and their pollen data are important for the diagnosis of the ancestral pollen features in eudicots. Our observations show these genera are defined by having microechinate microreticulate exine ornamentation, perforate tectum, columellate morphology of the infratectum and the existence of a foot layer and endexine. The presence of lamellations is detected during the early stages of development in the nexine of both genera, especially in the apertures. Euptelea presents remains of the primexine layer during the whole maturation process, a very thin foot layer, and a laminate exinous oncus in the apertural region formed by ectexine and endexine elements. Pteridophyllum has a thicker tectum than Euptelea, a continuous foot layer and a thicker endexine. In the apertures, the exinous oncus is formed by islets and granules of endexine, in contrast to the Euptelea apertures. The secretory tapetum produces orbicules in both genera, but they have different morphology and electron-density. Comparisons with pollen data from related orders and families confirm the ancestral states for the pollen of eudicots proposed in previous studies: reticulate and echinate surfaces, columellate infractectum and a thin foot layer relative to the thickness of the ectexine. According to our observations, we propose considering the possibility of a polymorphic state for the aperture number in the ancestor of Ranunculales, and suggest the development of orbicules as the ancestral state in this order.     

A unique pollen wall mutation in the family Compositae: ultrastructure and genetics

During a routine screening of pollen fertility in the n = 2 chromosome race of Haplopappus gracilis, a spineless pollen wall mutation was discovered that renders the otherwise functional pollen grains completely unrecognizable as Compositae pollen. Normal Haplopappus pollen is characterized by an outer layer, the ektexine, consisting of large spines supported by a roof (tectum), which in turn is supported by collumellae that are joined basally. A large cavity (cavea) stretches from aperture to aperture and separates columellae bases from the final ektexine unit, the foot layer. The spines, tectum, columellae, and columellae bases are filled with perforations (internal foramina), while the foot layer is without them. Immediately underlying the foot layer is a thickened, lamellate, disrupted, internal foramina-free second exine layer, the endexine. In contrast, the mutant pollen ektexine is a jumble of components with randomly dispersed spines as the only clearly definable unit. The endexine layer is similar to the endexine in normal pollen. The mutation apparently disrupts only the organization of ektexine units, and mutant pollen appears to be without the caveae and foot layer characteristic of normal pollen. In genetic tests, the mutant allele is recessive. There is a simple Mendelian pattern of inheritance of the mutant gene, and its phenotype is under sporophytic control.     

Comparative pollen morphology of Glechoma and Marmoritis (Nepetinae,Lamiaceae)

Abstract The pollen morphology of 13 taxa (34 specimens) of the genera Glechoma L., and Marmoritis Benth. was investigated in detail using light, scanning electron, and transmission electron microscopy. Pollen grains of all studied taxa are small to large in size (P= 32.5–60.4 μm, E= 20.2–50.5 μm), prolate‐spheroidal to prolate in shape and mostly hexacolpate (the amb more or less circular or rarely ellipsoid) with granular membranes. The sexine ornamentation of Glechoma is bireticulate; the muri of the primary reticulum are irregularly circled, and lumen size is short. In contrast, the sexine surfaces of the Marmoritis pollen tend to more elongate or wider at the muri of the primary reticulum than those of the Glechoma. The pollen wall stratification of selected taxa (three from Glechoma and one from Marmoritis) is characterized by unbranched columellae, and continuous or distinctly discontinuous endexine based on transmission electron microscopy observation. The results of Glechoma and Marmoritis reveal rather similar pollen morphological features, however, fine details of sexine ornamentation are characteristic to differentiate the pollen taxa. Although these differences may be useful in establishing the taxonomic boundary between two genera, they are too weak to segregate diagnostic characters.     

POLLEN APERTURE EVOLUTION AMONG THE SUBFAMILIES PERSOONIOIDEAE,SPHALMIOIDEAE, AND CARNARVONIOIDEAE (PROTEACEAE)

Pollen apertures were analyzed among the subfamilies Persoonioideae (seven genera; ca. 95 spp.), Sphalmioideae (one genus; one spp.), and Camarvonioideae (one genus; two spp.). Pollen was examined by light microscopy, cryosection, and transmission electron microscopy. Completed studies of pollen apertures among Grevilleoideae (ca. 40 genera; ca. 800 spp.), one of two major subfamilies in Proteaceae, provide a basis for comparison and analysis of aperture evolution among these subfamilies. Aperture characters within Persoonioideae are unique among Proteaceae examined to date. Five distinct aperture types occur among the three subfamilies, three of which (Placospermum, Persoonia, Bellendena) are restricted to Persoonioideae. Sphalmioideae and Camarvonioideae each exhibit a unique aperture organization. The most primitive aperture organization, and one unique to Placospermum, exhibits three main features: 1) a thin, granular endexine continuous around the grain; 2) a heterogeneous foot layer throughout the grain with increased disruptions at the aperture; and 3) only slight differences in exine characters between apertural and nonapertural regions. The Persoonia aperture type represents the next stage of aperture evolution which involves loss of endexine, restriction of a heterogeneous foot layer to the aperture, and marked differences in exine characters between apertural and nonapertural regions. The uniformly homogeneous ektexine in both nonapertural and aperture regions in Bellendena has developed independently. Sphalmium exhibits a primitively thin granular endexine though the restriction of endexine to the aperture is a derived condition. Carnarvonia exhibits several pollen characters also found among Grevilleoideae including: 1) a homogeneous nonapertural ektexine; 2) a slightly heterogeneous apertural ektexine; 3) a lamellate/granulate endexine organized into irregularly shaped “clumps” clustered around the aperture; and 4) a clear demarcation between apertural and nonapertural exine. These characters support the hypothesis that Carnarvonia may have diverged early from the pre-Grevilleoids.     

Change of Floral Orientation within an Inflorescence Affects Pollinator Behavior and Pollination Efficiency in a Bee-Pollinated Plant,Corydalis sheareri

Vertical raceme or spike inflorescences that are bee-pollinated tend to present their flowers horizontally. Horizontal presentation of flowers is hypothesized to enhance pollinator recognition and pollination precision, and it may also ensure greater consistency of pollinator movement on inflorescences. We tested the hypotheses using bee-pollinated Corydalis sheareri which has erect inflorescences consisting of flowers with horizontal orientation. We altered the orientation of individual flowers and prepared three types of inflorescences: (i) unmanipulated inflorescences with horizontal-facing flowers, (ii) inflorescences with flowers turned upward, and (iii) inflorescences with flowers turned downward. We compared number of inflorescences approached and visited, number of successive probes within an inflorescence, the direction percentage of vertical movement on inflorescences, efficiency of pollen removal and seed production per inflorescence. Deviation from horizontal orientation decreased both approaches and visits by leafcutter bees and bumble bees to inflorescences. Changes in floral orientation increased the proportion of downward movements by leafcutter bees and decreased the consistency of pollinator movement on inflorescences. In addition, pollen removal per visit and seed production per inflorescence also declined with changes of floral orientation. In conclusion, floral orientation seems more or less optimal as regards bee behavior and pollen transfer for Corydalis sheareri. A horizontal orientation may be under selection of pollinators and co-adapt with other aspects of the inflorescence and floral traits.     

Pollen morphology of Chinese Curcuma L. and Boesenbergia Kuntz (Zingiberaceae): Taxonomic implications

In order to find new non-molecular evidence to support the phylogenetic and taxonomic position, pollen grains of 20 populations of 16 species of Chinese Curcuma L. and Boesenbergia Kuntz (Zingiberaceae) were investigated under SEM and TEM. The pollen grains are spherical and ovoid, nonaperturate. The pollen wall is composed of a very thin exine and a thick intine. The exine is psilate or echinate. The intine consists of two layers, i.e., a thick, channeled layer (exintine) and an inner homogenous layer (endintine). The results reveal morphological congruence between the pollen grains of species of Curcuma, which according to DNA sequence data appears to be a polyphyletic genus. However the uniform pollen morphology in Curcuma provides no evidence to divide this genus into separate taxonomic entities. Our results on pollen morphology also do not provide any additional evidence to either unite or segregate Boesenbergia albomaculata and Curcumorpha longiflora in the same genus and demonstrate that more taxonomic data on the genus Boesenbergia and its relatives are needed before a final decision can be made.     

Causes of Low Fruit and Seed Set in Bird-Pollinated Anagyris foetida (Leguminosae): Pollen Limitation and Other Extrinsic Factors

We studied the extrinsic factors behind the low number of offspring in two populations of Anagyris foetida, a bird-pollinated leguminous shrub of the Mediterranean region. Fruit initiation was pollen limited, but fruit maturation was not. This limiting effect varied between flowering seasons and between populations, and also within a given phenological stage. In the first season, the early flowers had the lowest fruit set, while in the second season fruit set was highest in these early flowers. This was possibly related to the pollination environment. Seed initiation (fertilized ovules) increased following pollen supplementation, but this was not translated into a significant increase in either the number of seeds per fruit, or seed mass. This indicates that seed initiation is pollen limited but that other factors (e.g., resource availability) are involved in seed maturation. Abiotic factors such as excess humidity during flowering were responsible for the loss of inflorescences, especially in one of the two populations. In this population, the prevailing wind in autumn-winter was less effective in drying the flowers when there was excess humidity. Also, significantly fewer inflorescences were lost from solitary plants than from clustered plants, probably reflecting the beneficial action of the wind and the greater light levels during flowering. Of the biotic factors analyzed, sheep predation was the most important, being worse in drought years. This predation, by affecting population density, could modify the plant-pollinator interaction and severely reduce the plant’s breeding success because of its pollen limitation.     

Pollen Organs and Seeds with Eucommiidites Pollen

The characteristic Mesozoic pollen genus Eucommiidites is described from pollen organs and seeds recovered in Cretaceous strata of North America and Europe. The pollen organs are from the lowermost Upper Cretaceous (Cenomanian) of Texas and are referred to Erdtmanitheca texensis gen. et sp. nov. They are spherical heads, composed of numerous, densely crowded, radiating pollen sacs that contain abundant well-preserved pollen. Combined LM, SEM and TEM investigations show that the pollen grains each have a distinct distal colpus flanked by two lateral colpi in an equatorial position. Pollen wall ultrastructure is gymnospermous with a thick lamellate inner layer (endexine) and an outer layer (ektexine) composed of a granular inner part and a homogeneous outer part. The endexine is thickened in the region of the colpi. Small seeds from the Lower Cretaceous (upper Berriasian to Valanginian) of Bornholm, Denmark contain abundant Eucommiidites pollen in their micropyles. The seeds are referred to Erdtmanispermum balticum gen. et sp. nov. They are ovoid, and weakly triangular in transverse section and gradually taper at the apex into an elongated tube. The megaspore membrane is granular and well developed, and apparently surrounded by three separate tissues interpreted as nucellus, a thin inner integument and a sclerified outer envelope. Eucommiidites pollen in the micropyles of the seeds has a laminated endexine and an ektexine comprising two homogeneous parts separated by a granular layer. Réévaluation of other seeds known to contain Eucommiidites pollen indicates that they share significant similarities with Erdlmanispermum and that they may have been produced by closely related plants. Comparison of “Eucommiidites plants” with other seed plants suggests that they are probably most closely related to the anthophytes comprising Bennettitales, Pentoxylales, Gnetales and angiosperms.     

Pollen morphology,ultrastructure and taphonomy of the Neuradaceae with special reference to Neurada procumbens L. and Grielum humifusum E.Mey. ex Harv. et Sond.

Pollen morphology and sporoderm ultrastructure of modern Neurada procumbens L. and Grielum humifusum E.Mey. ex Harv. et Sond. were studied using light (LM) and electron (SEM and TEM) microscopy. Additionally late Holocene pollen of the Grielum-type was studied using LM. Systematic and ecological aspects have been discussed for the family Neuradaceae. The pollen grains of the studied species are characterized by similarities in size, shape, aperture type and differences in exine sculpture (reticulate semitectate exine in Neurada and finely reticulate to foveolate in Grielum) and sporoderm ultrastructure. The cavea in the exine is situated between the ectexine and endexine which are connected near the aperture region only. A combination of the palynological characters of the Neuradaceae (semitectate exine, rather loose columellae, interrupted foot layer, the cavea in the exine) increases the pollen plasticity, allowing considerable changes of the pollen grain volume but still remains insufficient to survive sharp fluctuations in hydration level.     

Variations of floral traits among different life forms illustrate the evolution of pollination systems in Potamogeton species from China

Four leaf morpha groups (floating-leaved, submerged broad-leaved, submerged linear-leaved and filiform-leaved) and three inflorescences type groups (erect, facultative floating and floating) can be found in the genus Potamogeton. Life forms in the genus correlate with pollination types (anemophily, epihydrophily and hydroautogamy). Floral traits vary greatly among different life forms and pollination types. Both floating and submerged broad-leaved species produce erect inflorescences that are suited for anemophilous pollination. These flowers also have the highest pollen/ovule (P/O) ratio (25,013) and the smallest pollen grain volume (6046 μm³). On the contrary, submerged filiform-leaved species produce only floating inflorescences and their flowers produce the largest pollen grains (17,567 μm³) and have the lowest P/O ratio (6752), which is typical of the syndromes of hydroautogamy and epihydrophily. The relatively prolific seed production (about 70%) and wide dispersal in the floating and submerged broad-leaved species and the submerged filliform-leaved species indicate that these two groups are well adapted to the aquatic environment. This is in contrast with the less well-adapted complex and unstable pollination system found in linear-leaved species. We propose that the pollination systems and life forms in the Potamogeton, have undergone modification in response to the demands imposed by the aquatic environment during the evolutionary process in the genus.