Demography, competitive interactions and grazing effects of intertidal limpets in southern New Zealand

Population dynamics and the effects of intraspecific competition on limpet growth and maintenance of bare patches were investigated for the intertidal limpet Cellana ornata (Dillwyn) at a boulder-dominated site and on a rocky platform near Kaikoura (South Island), New Zealand. Distribution and abundance patterns of C. ornata were described in relation to other biota and tidal height. C. ornata occurred almost exclusively in patches devoid of macroalgae, particularly in the mid-tidal zone. Both adult and juvenile limpets were most abundant on the tops of boulders, where their numbers were positively correlated with barnacle cover, which averaged 77%. The size structure and growth patterns of C. ornata were different between populations. Mark-recapture studies showed that the slopes of annual growth increments regressed on initial sizes were similar at both sites but that the annual increments on the platform were about 6 mm greater than on boulders. Growth virtually ceased at 27 mm for limpets at the barnacle-dominated boulder site and at 40 mm at the platform site. Recruiting cohorts had 20% survival on boulders and 37% on the platform during their first year. The largest size classes at both sites had around 57% annual survival. To test the effects of varying limpet densities on the growth and mortality of limpets and the maintenance of bare patches, densities of C. ornata were experimentally increased at both sites. Beyond a density of 4 per 0.25 m², sizes and survival of limpets were reduced at both sites, but the effect was more pronounced at the boulder site. Limpets at the boulder site were more effective at maintaining bare space than those on the reef platform. Enclosing limpets in plots with and without barnacles showed that C. ornata and a co-occurring species (Cellana denticulata (Martyn)) grazed more effectively and had greater growth in cleared plots. Overall, there was considerable variation in the demographics of C. ornata between populations driven by site-specific characteristics.     

Sexual dimorphism of body size and shell shape in European tortoises

Adult body size and shape were examined in almost 1400 individuals of the tortoises Testudo graeca , T. hermanni and T. marginata from Greece. The size at maturity was greater in females than in males in all three species. Maximum and mean adult sizes were also greater in females than in males in T. graeca and T. hermanni . Males grew to a larger size than females in T. marginata , and mean adult size was similar in the sexes in this species. Sexual dimorphism of shape (adjusted for size covariate) was shown in most of the characters examined, and the degree of this dimorphism differed significantly among the three species. Differences were related to their contrasting courtship behaviours: horizontal head movements and severe biting in T. marginata , vertical head bobs and carapace butting in T. graeca , and mounting and tail thrusting in T. hermanni . There was no difference in the frequency of observations of courtship or fighting among the three species, but courtship was about 10 times more common than combat in males. All species showed greatest courtship activity in autumn; copulation was rarely observed in T. hermanni (only 0.36% of courting males) and not seen in the other species in the field. Observations made throughout the activity season indicated that feeding was equally common in males and females in all three species. Differences in shape were more likely to be the result of sexual selection than of natural selection for fecundity. Detailed predictions are made for sexual dimorphism of other characters in these species.     

Analysis of inter- and intra-specific competition amongst intertidal limpets with different methods of feeding

Summary The prosobranch limpet Cellana tramoserica grazes on microalgae, including the spores of macroalgae, and coexists with the pulmonate limpets Siphonaria denticulata and S. virgulata at mid-tidal levels on sandstone shores in New South Wales. These siphonarians graze on macroalgae, leaving the basal parts of the thalli intact. Where Cellana graze, they are capable of removing all algae; where Siphonaria graze, they leave at least a thin film of alga on the rocks, which is available to Cellana. S. denticulata normally show invariant homing behaviour, whereas S. virgulata tend to move around at random when Cellana are present, but apparently home when Cellana are absent. Both siphonarians have been observed to show opportunistic behavioural responses by moving towards patches of macroalgal foods when they become available.Experimental ecclosures of limpets at different densities and in different combinations revealed that Cellana tramoserica suffered increased mortality and reduced growth due to intraspecific competition when at increased densities. There was no effect on Cellana of increased densities of either species of Siphonaria. Nor was there any interspecific interaction between the siphonarians. Both species of Siphonaria showed some reduction of growth at increased intraspecific density. More importantly, both showed increased mortality when enclosed with low densities of Cellana. Larger densities of Cellana had no effect; the numbers of Cellana could not be maintained because of the reductions caused by intraspecific competition. Even after 27 weeks in enclosures with Cellana, the numbers of Siphonaria never declined to zero in any experimental enclosure. Thus, Cellana has a competitive effect on the survival of siphonarian limpets, but is unable to exclude them from an area of the shore. Siphonaria spp., in contrast, have no effect on Cellana.The nature of the competitive interactions between these types of limpets is explaied in terms of their methods of feeding; Cellana can exploit the food-resource before it reaches a suitable size for Siphonaria. The coexistence of Siphonaria spp. with Cellana is discussed with respect to the behaviour of the pulmonates. Intraspecific competition leading to reduced densitities of Cellana, however, will ensure that Cellana cannot exploit all the food resources, and some will be available to Siphonaria. The consequences of inter- and intra-specific competition among grazing gastropods are discussed with reference to the structure of intertidal communities, and it appears that competition for food is fundamentally different from competitive interactions for space in the organization of such communities.     

Effects of shape,size, and pyrene doping on electronic properties of graphene nanoflakes

Effects of size, shape, and pyrene doping on electronic properties of graphene nanoflakes (GNFs) were theoretically investigated using density functional theory method with PBE, B3PW91, and M06-2X functionals and cc-pVDZ basis set. Two shapes of zigzag GNFs, hexagonal (HGN) and rhomboidal (RGN), were considered. The energy band gap of GNF depends on shape and decreases with size. The HGN has larger band gap energy (1.23–3.96 eV) than the RGN (0.13–2.12 eV). The doping of pyrene and pyrene derivatives on both HGN and RGN was also studied. The adsorption energy of pyrene and pyrene derivatives on GNF does not depend on the shape of GNFs with energies between 21 and 27 kcal mol^?1. The substituent on pyrene enhances the binding to GNF but the strength does not depend on electron withdrawing or donating capability. The doping by pyrene and pyrene derivatives also shifts the HOMO and LUMO energies of GNFs. Both positive (destabilizing) and negative (stabilizing) shifts on HOMO and LUMO of GNFs were seen. The direction and magnitude of the shift do not follow the electron withdrawing and donating capability of pyrene substituents. However, only a slight shift was observed for doped RGN. A shift of 0.19 eV was noticed for HOMO of HGN doped with 1-aminopyrene (pyNH₂) and of 0.04 eV for LUMO of HGN doped with 1-pyrenecarboxylic acid (pyCOOH).

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Graphical Abstract HOMO and LUMO Energies of pyrene/pyrene derivatives doped Graphene Nanoflakes

     

Effects of monovalent cations on red cell shape and size

Human erythrocytes were incubated in isotonic solutions of different monovalent cations. The apparent size of the red cells measured on scanning electron microscopic pictures decreases in the order Li+ greater than Na+ = K+ greater than Rb+. These differences in size are abolished after pretreatment with trypsin, which removes a large part of the charges associated with membrane glycoproteins. Shape alterations are also observed. Normal biconcave shapes are visible after Na+ or K+ incubation, whereas Li+ leads to flabby, flattened cells with a certain tendency to crenation, and Rb+ causes more pronounced biconcavity with a certain tendency to cupping. The overall effects of pretreatment with trypsin are similar to those of Li+. Our results provide evidence that the electrostatic repulsion of glycoproteins and other charged membrane components may play an essential role in maintaining red cell shape.     

Effects of monovalent cations on red cell shape and size

Human erythrocytes were incubated in isotonic solutions of different monovalent cations. The apparent size of the red cells measured on scanning electron microscopic pictures decreases in the order Li⁺>Na⁺=K⁺>Rb⁺. These differences in size are abolished after pretreatment with trypsin, which removes a large part of the charges associated with membrane glycoproteins. Shape alterations are also observed. Normal biconcave shapes are visible after Na⁺ or K⁺ incubation, whereas Li⁺ leads to flabby, flattened cells with a certain tendency to crenation, and Rb⁺ causes more pronounced biconcavity with a certain tendency to cupping. The overall effects of pretreatment with trypsin are similar to those of Li⁺. Our results provide evidence that the electrostatic repulsion of glycoproteins and other charged membrane components may play an essential role in maintaining red cell shape.     

Effects of patch size and position above the substratum during early succession of subtidal soft-bottom communities

Early macrobenthos succession in small, disturbed patches on subtidal soft bottoms is facilitated by the arrival of post-larval colonizers, in particular by active and passive dispersers along the seafloor or through the water column. Using a field experiment at two contrasting sites (protected vs. exposed to wave action), we evaluated the role of (a) active and passive dispersal through the water column and (b) the influence of small-scale spatial variability during succession of subtidal macrobenthic communities in northern Chile. Containers of two sizes (surface area: small—0.12?m² and large—0.28?m²) at two positions above the natural substratum (height: low—3?cm and high—26?cm) were filled with defaunated sediment, installed at two sandy sublittoral sites (7–9?m water depth) and sampled after 7, 15, 30, 60 and 90?days, together with the natural bottom sediment. The experiment took place during austral fall (from late March to early July 2010), when both larval and post-larval stages are abundant. At the exposed site, early succession was driven by similar proportions of active and passive dispersers. A sequence from early, late and reference communities was also evident, but container position and size affected the proportional abundance of dispersal types. At the protected site, the successional process started with abundant colonization of active dispersers, but toward the end of the experiment, the proportion of swimmer/crawlers increased, thus resembling the dispersal types found in the natural community. At this site, the position above the sediment affected the proportional abundance of dispersal types, but patch size had no effect. This study highlights that macrobenthic post-larvae can reach at least 26?cm high above the bottom (actively or passively, depending on site exposure), thus playing an important role during early succession of sublittoral soft bottoms. The active or passive use of the sediment–water interphase may also play an important role in the connectivity of benthic populations and in the recovery after large-scale disturbances of sublittoral habitats.     

Effects of body size and water temperature on grazing rates of four intertidal gastropods

Abstract Rates of radular activity were measured in four species of Australian gastropods. The species were Austrocochlea constricta, Bembicium nanum, Cellana tramoserica, and Nerita atramentosa. Work was done in Botany Bay at Cape Banks, NSW, Australia from September 1989 to June 1990. A hydrophone was used to record radular raspings of snails in situ. Averages based on all observations showed Cellana had the fastest rate with 80 rasps/min, and was followed by Austrocochlea (71 rasps/min), Bembicium (57), and Nerita (39). Faster rates were associated with warmer water temperature and smaller body sizes in all species except Bembicium. The potential for these differences in radular activity affecting competitive interactions is discussed.     

The fibrinogen molecule: its size,shape, and mode of polymerization

Improved electron micrographs of the shadow-cast bovine fibrinogen molecule have been obtained establishing its general morphology and dimensions in the dry state. It consists of a linear array of 3 nodules held together by a very thin thread which is estimated to have a diameter of from 8 to 15 A, though it is not clearly resolved. The two end nodules are alike but the center one is slightly smaller. Measurements of shadow lengths indicate that nodule diameters are in the range 50 to 70 A. The length of the dried molecule is 475 +/- 25 A. Adopting the molecular volume from previous physical chemical data and the general morphological features and length from electron microscopy, we calculate the diameters of the end nodules to be 65 A and the center one as 50 A. The model of the molecule so obtained is consistent with the electron microscopical observations and the data from physical chemistry. The intermediate polymers formed when fibrinogen is activated with thrombin were also examined and found to be end-to-end aggregates of altered fibrinogen molecules which shrink in length during the process. Intermediate polymer lengths are from 1000 to 5000 A. The nodular nature of fibrinogen, its shrinkage and end-to-end aggregation on polymerization permits us to deduce an explanation for the system of cross-bands previously observed in stained fibrin fibrils.     

Effects of X chromosome on size and shape of body: an anthropometric investigation in 47,XXY males

The effects of an extra X chromosome on size and shape of body and head were studied in 47,XXY males; 25 anthropometric measurements were recorded from 29 adult 47,XXY males and compared with those of male relatives and control males. In stature, arm length, leg length, triceps skinfold, and subscapular skinfold 47,XXY males were larger and in biacromial diameter, bideltoid breadth, wrist breadth, and in most head dimensions smaller than normal males. Arm length was increased less than leg length. Increase in stature seemed to be caused solely by increased leg length, and the somewhat feminine proportions in trunk were caused by decrease in biacromial diameter. Correlations of the body and head dimensions between 47,XXY males and their male relatives were found to be normal. The present findings support the earlier proposals that X chromosome carries genes which influence linear growth. It is suggested that the reduction in biacromial diameter is caused by lowered plasma testosterone level which may also have affected sitting height. The control of body and head dimensions seems to be maintained relatively normal.     

Effects of temperature on the size and shape of Escherichia coli cells

Two substrains of Escherichia coli B/r were grown to steady-state in batch cultures at temperatures between 22 and 42° C in different growth media. The size and shape of the cells were measured from light and electron micrographs and with the Coulter channelizer. The results indicate that cells are shorter and somewhat thicker at the lower temperatures, especially in rich growth media; cell volume is then slightly smaller. A lower temperature was further found to increase the relative duration of the constriction period. The shapes of the cell size distributions are indistinguishable, indicating that the pattern of growth of the cells is the same at all temperatures. The adaptation of the cells to a temperature shift lasted several generations, indicating that the morphological effects of temperature are mediated by the cell's physiology.     

Light and electron microscopical investigations on shell pores (caeca) of fissurellid limpets (Mollusca: Archaeogastropoda)

The organic content (caecum) of the shell pores of five species of fissurellid (key-hole) limpets was investigated by means of light and electron microscopy. Fissurellid caeca are unbranched, polycellular extensions of the mantle epithelium, which penetrate the whole shell and contact the periostracum via organic fibres called 'brush'. They are built up by two cell-types, which enclose a central lumen, and are formed concurrently with the shell itself. Because of fundamental structural and developmental differences, the superficially similar shell pores of bivalves (caeca) and polyplacophorans (aesthetes) are considered as analogous rather than homologous organs. Similarities are found with brachiopod caeca, probably owing to similar functions.     

Age and growth rate determinations of an intertidal bivalve, Phacosoma japonicum, using internal shell increments

A venerid bivalve Phacosoma japonicum (Reeve) occurring commonly in the Japanese coastal area preserves periodic growth lines in the shell cross-section. Long-term shell growth patterns of this species have been traced for many individuals on the intertidal flat of the Seto Inland Sea, west Japan. Sclerochronological analysis of these individuals and specimens collected monthly shows that several growth cessation marks within their shells are formed during the winter of each year prior to spawning. Hence the marks were used for age and growth rate determinations. As large individuals showed little shell growth for more than two years after the formation of 7 or 8 annual increments, this species probably has a lifespan of more than ten years. Shell growth patterns of this species based on annual increments can be accurately approximated by a von Bertalanffy curve. The number of microgrowth increments formed during a year tends to decrease with age, although it varies markedly among specimens of the same age. Furthermore, even in summer during rapid shell growth, the microgrowth increments do not represent daily and/or sub-daily tidal rhythms in many specimens. The results of this study and those by several authors strongly suggest that the annual increments are the key for age and growth rate determinations of both living and fossil bivalve species.     

Measuring dimensions: the regulation of size and shape

Over many years evidence has accumulated that plants and animals can regulate growth with reference to overall size rather than cell number. Thus, organs and organisms grow until they reach their characteristic size and shape and then they stop - they can even compensate for experimental manipulations that change, over several fold, cell number or average cell size. If the cell size is altered, the organism responds with a change in cell number and vice versa. We look at the Drosophila wing in more detail: here, both extracellular and intracellular regulators have been identified that link cell growth, division and cell survival to final organ size. We discuss a hypothesis that the local steepness of a morphogen gradient is a measure of length in one axis, a measure that is used to determine whether there will be net growth or not.     

Regularity of species richness relationships to patch size and shape

This study aims to assess the degree of regularity in the effect of patch size and patch shape on plant species richness across a macroscale region, and to evaluate the implications for nature conservation.
Our study area covers south-eastern Norway and contains 16 agricultural landscapes with 2162 patches.
To analyse regularity a local linear mixed model (LLMM) was applied. This procedure estimates the richness trends due to shared effects of size and shape, and simultaneously provides the landscape-specific random effect. The latter is a direct estimate of the degree of irregularity between the landscapes, conditioned on specific values of size and shape.
The results show a positive interaction between the shape and size of patches, which is repeated for all landscapes. The shape of the patches produces more regular patterns in species richness than the size of patches. This we attribute to effects of dispersal and distance to neighbouring patches of different environmentally conditioned species pools. Large and complex patches have shorter average distance to neighbouring patches (of different types) than large simple-shaped (circular) patches have. We attribute the higher species richness of the former, given a similar area, to a higher number of species dispersed from the outside into the more complex plot. For small patches, however, the distance to the edge is short relative to normal dispersal distances, for patches of all shapes. This explains why the positive effect of shape complexity on species richness is stronger for large patches. This interpretation is supported by a strong spatial correlation conditioned on the most complex patches.
Theories of dynamics in biodiversity in patchy landscapes must consider shape as a regulator at the same level as size, and both shape and size of patches should be simultaneously taken into account for management planning.