The functional significance of aperture form in gastropods

Aperture form of marine prosobranch gastropods has evolved under the influence of a number of different selective forces, including: generation of shell form; protection from predation; accommodation of the foot during clamping behavior: and accommodation of water currents in and out of the mantle cavity. Aperture form correlates positively with foot shape in most gastropods and foot shape, in turn, correlates moderately well with substrate preference. Almost all gastropods that have non-round apertures elongate the aperture parallel to the foot so that water currenth tend to flow anteriorly to posteriorly. Fresh-water pulmonates have responded to somewhat different stresses. They exhibit clamping behavior and thus show correspondence between foot shape and aperture shape. They show less apertural strengthening as crab (or crayfish) predation is less of a factor and presumably because calcium carbonate is less available. They also lack anterior-posterior apertural elongation due to the absence of water currents through their mantle cavity. Due to the absence of mantle cavity water currents and clamping behavior, terrestrial gastropods do not show the apertural modifications associated with these two factors. In addition. few adaptations of apertural form are present to resist predation. Instead, many of the apertural modifications of terrestrial pulmonates seem to be concer-ned with the problems of water loss during estivation.     

Multilamellar bodies linked to two active plasmalemma regions in the pollen grains of Sarcocapnos pulcherrima

The presence of visible multilamellar bodies in the cytoplasm of pollen grains of at least seven species of the family Papaveraceae has led us to study the behaviour of these bodies during pollen-grain ontogeny and in growing pollen tubes of Sarcocapnos pulcherrima C. Morales & R. Garcia germinated in vitro. Our transmission-electron-microscope (TEM) studies in pollen grains show that the multilamellar bodies may be classified as: 1) small, isolated and placed in the region of apertures in the cytoplasm; and 2) large, in clusters and in contact with the active plasmalemma apertures only when tubules are being formed in the apertural intine. Similar types of multilamellar bodies to those observed in the pollen apertures can be seen near the apex of the growing pollen tube (small and isolated) and in contact with the apex plasmalemma (large and clustered). Our results support the hypothesis that the multilamellar bodies are functionally linked to moments when the cytoplasmic membrane is very active. We have also linked the multilamellar bodies to Golgi vesicles as they both react positively to acid-phosphatase (AP) staining and also to the plasmalemma by the thiocarbohydrazide-silver proteinate-staining (TCH-Sp) electron-contrasting technique.     

The geographic, taxonomic and temporal distribution of determinate growth in marine gastropods

Many marine gastropods are characterized by determinate growth, as inferred from the presence of unique terminal elaborations of the shell's aperture. Although determinate growth has evolved repeatedly in most major gastropod clades, it is especially frequent among siphonate caenogastropods. Analyses of shallow-water assemblages show that the incidence of species with determinate growth is far higher in the tropics (especially the tropical Pacific and Indian Oceans) than at higher latitudes. Compilations of fossil assemblages from warm-water environments indicate that, although determinate growth occurred in some Palaeozoic gastropods, it became widespread only in the Neogene. In some groups, terminal apertural elaborations arose in lineages whose growth was more or less continuous and indeterminate, but in others it was derived either from or was ancestral to episodic growth. The hypothesis that periodic or terminal apertural elaborations evolved as a means to dispose of calcium carbonate once growth in the spiral direction ceased is rejected in favour of functional interpretations. Among the latter, the roles of modified apertures in defence and in mate recognition are explored, but no firm conclusion regarding the latter possibility can be drawn owing to our ignorance of mate recognition in gastropods.     

The Microfibrillar Component of the Pollen Intine Some Structural Features

The microfibrillar polysaccharide component of the pollen intinecan be isolated by progressive chemical digestion of the exineand the cellular contents and the extraction of the matrix materials.The resulting intine ‘ghosts’ reveal various characteristicstructural features. The microfibrils have apparent individualdiameters in the range of 5–15 nm, but they are commonlyassociated laterally to form ribbons, or aggregated in strandsor cables of dimensions great enough to be resolved with theoptical microscope. These often show preferred orientations,which can be associated with pollen grain shape and with thedisposition-of the germination apertures. The apertural intinemay be structurally complex, as in Abutilon hybridum, where,after the removal of the exine, the polysaccharide caps whichoverlie the protein storage sites of the pollen grain wall retainthe elaborate patterning of the original cytoplasmic evaginationsfrom the vegetative cell. Secale cereale, Narcissus pseudonarcissus, Abutilon hybridum, Crocus vernus, pollen grain, intine, exine, wall pattern, germination apertures, polysaccharide microfibrils, fluorescence microscopy     

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).     

Morphologie, fonction et évolution de quelques grains de pollen normapolles du Crétacé supérieur

Upper Cretaceous pollen grains with a thick wall (exine) and sometimes protruding apertures, included in the Normapolles Pflug, are considered from a functional point of view. Because of the high development of compression-resistance features correlative with a high protection of the gametophyte, an entomophilic phase is supposed in the dissemination of some types, but anemophily appears to have been a developmental trend of apertural specialization for others. As these functional features are characteristic of the outer part of the wall (the sexine), the tetrahedral morphology of the inner part (the nexine) and the lack of evidence that it influenced the whole adaptive system suggest that angiospermic constraints have acted first at a pteridophytic stage.     

A tubicolous animal from the Hunsrück Slate (Lower Devonian,Southern Germany)

A small tube-dwelling organism is interpreted as living in the sediment. This life habit is inferred from the difference between lined burrows and tubes and between soft and rigid tubes. Lamellar extensions of the body-wall may have provided irrigation for the vermiform part of the body. The gaping aperture lacks an operculum and may have been open permanently; the apertural rim was strengthened by spicules. The specimen lacks segmentation and is thus not an annelid worm. Considering the theoretical suitability of the crinoid “forest” in the Hunsrück sea for such worms, the absence of Annelida is curious.     

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.     

Remarks on the systematic position of Lithocodium Elliott, a problematic microorganism from the Mesozoic carbonate platforms of the Tethyan realm

The encrusting genus Lithocodium Elliott, widespread in Tethyan Triassic-Cretaceous carbonate platforms and interpreted in the past as a codiacean alga, is presently regarded as a colony of calcified cyanobacteria, a microorganism incertae sedis or a loftusiacean foraminifer. The absence of apertures connecting neighbouring cavities, as well as the very irregular form and arrangement of these hollows within a frequently massive micritic matrix, excludes assigning Lithocodium to the foraminifers, but could rather suggest colonies of calcified cyanobacteria.     

Spinihornera, a New Bryozoan Genus

Spinihornera is proposed as a new genus for the species "Hornera" spinigera Kirkpatrick, 1888, and is referred to the family Horneridae Smitt. This genus is characterized by its double walled, hornerid structure, and by having the autozooecial apertures arranged in lateral lines, pinnate branches and apertural spines.     

Pollen morphology of the tribes Eritrichieae and Cynoglosseae (Boraginaceae) in the Iberian Peninsula and its taxonomic significance

DÍEZ, M. J. & VALDÉS, B., 1991. Pollen morphology of the tribes Eritrichieae and Cynoglosseae (Boraginaceae) in the Iberian Peninsula and its taxonomic significance.
The pollen morphology of 33 species of Boraginaceae from the Iberian Peninsula belonging to the tribes Eritrichieae and Cynoglosseae has been studied by light and scanning electron microscopy. The results confirm the eurypalinous character of this family, in which a large number of species can be recognized by their pollen characters.
The pollen of these two tribes is basically heterocolpate, a type which is very uncommon in the angiosperms, with 6, 8 or 10 apertures, sometimes with ectoapertures anastomosed at the equator (ectocingulum), or diorate, minute or small in size, outline elliptic or rectangular-elliptic in equatorial view, with or without a constriction at the equator, exine 0.5-1.0 urn thick and the tectum psilate with granules around the colpi. The similarity in shape and apertural system found in several genera such as Asperugo, Gyrocarion and Omphalodes, and some species of Myosotis, does not permit the two tribes to be distinguished on pollen characters. The generic limits of Cynoglossum and the infrageneric classification of Myosotis are discussed.     

Mechanical constraints on aperture form in gastropods

Gastropod apertures reflect expanded states of their mantle edge under variable boundary conditions. The apertures are divided into two groups: apertures without distinct overlap zones (e.g., whorl overlap) and those with overlap zones. Each group follows a unique morphological rule. Apertures without overlap zones are generally circular in outline. Apertures with overlap zones are either elongated perpendicularly to the overlap zone or inflated abapically. Moreover, the latter abapically inflated apertures are generally accompanied by a straight section anterior to the overlap zone along the columellar axis (columellar part). Numerical analysis of an elastic double membrane tube whose main frame simulates the gastropod mantle indicates that these morphological rules are the products of mantle edge expansion under the condition that the head-foot mass presses against the mantle edge in both the overlap zone and the columellar part. The mantle edge in these two zones is thus in a completely or partly fixed boundary condition at the moment of shell growth. The rest of the mantle edge is free to expand either symmetrically or asymmetrically. It is hypothesized that the head-foot mass is a driving force for regulating the pattern of shell coiling and apertural shape.     

Divergent shell shape as an antipredator adaptation in tropical land snails

Although many land snails exhibit amazingly divergent shell shapes in the tropics, the functions of these remain obscure. Here we show that a modified aperture shape acts as an impediment specifically to predation by a snail-eating snake. Pareas iwasakii (Colubridae: Pareatinae) uses a unique method to feed on land snails: the snake extracts the soft body from the shell through the aperture by alternately retracting its mandibles. The snail Satsuma caliginosa (Camaenidae: Camaeninae) has apertural variation in regard to the presence of snail-eating snakes. Our experiments demonstrated that the distorted aperture mechanically impeded predation by this gape-limited predator, interrupting the mandibular movements. In contrast, congeneric snails with round apertures did not escape predation by snakes. The paleobiogeography of the focal area indicates that the subspecies Satsuma caliginosa picta, which does not have apertural modification, was derived from a defensive ancestor after the extinction of snail-eating snakes. Our study suggests a possibility that snail-eating snakes are responsible for divergent shell shapes in a variety of tropical land snails.     

A contribution to the ultrastructural knowledge of the pollen exine in subtribe Inulinae (Inuleae, Asteraceae)

To better understand the relationships within the Asteroideae and Inuleae, the structure of the pollen exine was investigated in seven genera and nine species of the subtribe Inulinae using LM, TEM and SEM. All taxa have a senecioid pattern of exine. The tectal complex consists of three main layers that differ in thickness and morphology: a tectum, a columellar layer, and a layer consisting of the basal region of the columellae. The absence or the vestigial condition of the foramina is considered as a plesiomorphy within the Asteroideae. All taxa have a complex apertural system that consists of an ecto-, a meso-, and an endoaperture. These apertures intersect respectively the tectal complex, the foot layer and the upper part of the endexine, and the inner layer of the endexine. A continuous transition among the different species of Inulinae was found for all the quantitative characters examined. This relative homogeneity of the pollen morphological characters enhances the naturality of the subtribe Inulinae.     

Release of proteins from the pollen wall of Linum grandiflorum

Summary The emission of proteins from the pollen wall of Linum grandiflorum stained with Coomassie blue was followed directly in moistened grains as well as in pollen prints. Within the first minute of the grain being moistened exine-borne proteins emerged from both inter-apertural and apertural sites; subsequently, proteins of a different nature were discharged from the apertures only. In a fraction of the grains the release of intine proteins was not preceded by that of exine proteins. Pin and thrum pollen did not differ in terms of mode or site of this protein emission. The presence and emergence of exine proteins from the apertures is explained by the process of infolding of the colpal wall at desiccation and its expansion at rehydration, which causes an initial trapping and subsequent re-exposure of surface materials. This explanation may also account for the occurrence of poral sporophytic proteins in the pollens of many dictoyledons.     

The Apertural Wall in Pollen of Linum grandiflorum

The apertural wall in tricolpate pollen of Linum grandiflorumwas investigated in order to understand its functioning duringdesiccation and rchydration. Whole and sectioned pollen grainswere studied with light or electron microscopy and by cytochemicalmeans. The areas of the apertures were examined in fresh drypollen, in grains moistened on agar gel or removed from compatiblestigmas, and in pollen from mature undehisced anthers The intine was found to consist of an inner ß-glucanlayer and an outer pectic layer. At the apertures the pecticlayer is thickened and overlaid by a ß-glucan layer.The pectinaceous intine stains red with basic fuchsin. The presenceof a third wall layer, the medine, was not confirmed. The aperturalintine thickenings possess considerable imbibitional capacityand at rehydration they appear as swollen lenticular bodies A procedure is described for obtaining intact exine free grains(EFG's) and whole, separated exines of L. grandiflorum. Invariably,the released EFG's consisted of protoplasts encased in the cellulosicintine. In most grains the outer intine remained attached tothe separated exine In L. grandiflorum the outer wall of the aperture expands whilethe protoplast and endintine are still infolded. Apparently,the exintine becomes detached from the endintine during desiccationand re-attaches at rehydration. It is suggested that the transientdetachment controls the influx of water into the vegetativecell Except for morph-specific exine processes no differences instructure of the aperture wall or its functioning at rehydrationwere observed between pin and thrum grains Pollen wallM, apertures, exintine, exine free grains, rehydration, desiccation, Linum grandiflorum     

Structure of the apertural sporoderm of pollen grains inEuphorbia andChamaesyce (Euphorbiaceae)

The tricolporate pollen grains of 38 Mediterranean and Macaronesian species ofEuphorbia L. andChamaesyce S. F. Gray have a special apertural sporoderm not found in the other taxa of theEuphorbiaceae. At the apertural margo the ectexine is thinner because of shorter columellae and the thin, fragmented or even absent foot-layer. Ectexinous granules, mixed with endexinous material, are present near the ora. The endexine is homogeneous and thickened under the colpi (at the end and at the proximity of the end of colpus). Around the ora, the endexine is granulate and lamellar with irregular cavities. The apertural intine presents a characteristic structure with thickenings running along both sides of the colpi. The arrangement and structure of these intinous thickenings depend on the distance from the ora. This special morphology of the intine is present in all taxa studied here. The genusEuphorbia is considered to be the most evolved taxon of this family. The characteristic apertural sporoderm may be an adaptative modification to different physiological conditions, so it may present an apertural mechanism which is more adapted to harmomegathic changes and thus facilitate the germination and the formation of the pollen tube.     

Distribution and abundance of the planktic foraminifer Neogloboquadrina pachyderma in sea ice of the Weddell Sea (Antarctica)

Summary Sea ice cores were obtained from eleven fast ice stations and one floe in the Weddell Sea, Antarctica in January–February 1985. All cores from the north eastern part of the Weddell Sea contained numerous living and dead planktic foraminifers of the species Neogloboquadrina pachyderma (Ehrenberg), while cores drilled in southern parts were barren of foraminifers with one exception. Foraminiferal abundances were variable, with numbers up to 320 individuals per liter melted sea ice. Distribution of foraminifers appears to be patchy, parallel cores taken less than 30 cm apart contained numbers which varied considerably. On the other hand, three cores taken on a transect each more than 3 km apart showed striking similarities. In general, small dead tests were found in the upper parts of the sea ice cores while large living individuals mainly occurred in lower sections. Abundant diatoms probably serve as a food source for the foraminifers. Correlation of foraminiferal abundance with salinity, chlorophyll and nutrient profiles are inconsistent. The possible mechanism of incorporation of N. pachyderma into the ice is discussed.