An arbitrary classification of the pore systems in avian eggshells

An arbitrary classification of the pore systems in shells has been based on the observations made during a survey with a scanning electron microscope of the eggs of c. 60 avian species. The classification is intended primarily to aid comparative studies of shell function and field studies of the overall contribution of the shell to the embryo's well-being. The location and form of the external orifice of the pore was used to define four major types of pore systems: (1) outer orifice open—the pore canal terminates at the surface of the shell; (2) outer orifice occluded—the pore canal terminates at the surface of the shell but the orifice is occluded by amorphous, waxy material; (3) outer orifice capped—the pore canal terminates at the outer edge of the true shell and its orifice is capped and, in many species, plugged by a stratum of vesicles, either organic or inorganic, which clothes the entire surface of the shell, and (4) outer orifice reticulate—the pore canal terminates at the edge of the column layer and its orifice is covered by a calcareous stratum containing a plexus of tubules. Each group was further divided on the form of the pore canals; shells having unbranched pore canals were put into one category and those in which some pores branched into another.     

The fine structure of the outer surface of the incubated eggshells of the Helmeted guinea fowl (Numidia meleagris)

The surface of the eggshells of the Helmeted guinea fowl (Numidia meleagris) was polished during incubation by the parent. Examination with the light microscope showed that the cuticle had been removed from the ridges on the outer surface of the shell and that the plugs in the outer orifice of the pore canals had acquired extraneous materials including grease. Studies with a scanning electron microscope revealed that the spheres that made up the pore plugs retained their identity even though they were stained. It was concluded that ridges on the shell surface protected the pore plugs from damage by attrition and that the plugs acted as filters thereby preventing nest debris from occluding the pore canals or contaminating the shell membranes.     

Structure of the shell and tertiary membranes of eggs of softshell turtles (Trionyx spiniferus)

Eggs of the turtle Trionyx spiniferus are rigid, calcareous spheres averaging 2.5 cm in diameter. The eggshell is morphologically very similar to avian eggshells. The outer crystalline layer is composed of roughly columnar aggregates, or shell units, of calcium carbonate in the aragonite form. Each shell unit tapers to a somewhat conical tip at its base. Interior to the crystalline layer are two tertiary egg membranes: the outer shell membrane and the inner shell membrane. The outer shell membrane is firmly attached to the inner surface of the shell, and the two membranes are in contact except at the air cell, where the inner shell membrane separates from the outer shell membrane. Both membranes are multi-layered, with the inner shell membrane exhibiting a more fibrous structure than the outer shell membrane. Numerous pores are found in the eggshell, and these generally occur at the intersection of four or more shell units.     

Aspects of shell formation in eggs of the tuatara,Sphenodon punctatus

The flexible shell from eggs of the tuatara (Sphenodon punctatus) is comprised of both calcareous and fibrous components. The calcareous material is organized into columns that extend deep into the fibrous shell membrane. Many of the fibers of the membrane are enclosed within the crystalline matrix of the columns. Columns widen and flatten slightly at the outer surface of the eggshell to form cap-like structures composed of a compact crystalline matrix containing no fibers. The outer surface of eggs laid prior to completion of shell formation consists of a series of nodes obscured by a densely fibrous matrix. Similar nodes also are found at the inner surface of partially shelled eggs. The nodes represent the outer and inner aspects of columns that had not completed formation prior to oviposition. Our interpretation is that a layer (or layers) of the shell membrane forms first, with nucleation of columns occurring shortly thereafter. Columns grow into the membrane a short distance and enclose fibers of the membrane, but the primary direction of column growth is toward what will become the outer aspect of the shell. Calcareous columns and the shell membrane form more or less in concert until crystal growth outstrips that of the membrane and a cap-like apex of compact crystalline material is formed. The end result is an eggshell in which the shell membrane and calcareous material form a single unit for much of the thickness of the shell.     

PROPERTIES OF AVIAN EGG SHELLS AND THEIR ADAPTIVE VALUE

I. An arbitrary classification of avian eggshells is proposed.
2. The role of the eggshell in conserving the water in eggs at oviposition is discussed. There is as yet no correlation between this property and the pore systems in avian eggshells.
3. The pore systems may act as diffusion pathways and the hypothesis has been advanced that in many eggs the shells are adapted so that restriction of gaseous diffusion by mud, preening oils and nest debris is prevented.
4. The mechanical properties of the shell are considered in the novel context of defence against (a) attrition that could lead to the pores being blocked with dust, and (b) cracking that would destroy the diffusion pathways noted in 3.
5. The overall objective of the review was to discuss the concept that avian eggshell are adapted to fit an egg to the nest environment.     

Differentiation and Growth of the Brachiopod Mantle

Cell differentiation in the mantle edge of Notosaria, Thecidelhnaand Glottidia, representing respectively, the impunctate andpunctate calcareous articulate and chitinophosphatic inarticulatebrachiopods, is described. Comparison of electron micrographssuggests that outer epithelium which secretes periostracum andmineral shell, is separated from inner epithelium by a bandof "lobate" cells, of variable width, exuding an impersistentmucopolysaccharide film or pellicle. The lobate cells alwaysoccupy the same relative position on the inner surface of theouter mantle lobe; but the outer epithelium is commonly connectedwith the inner surface of the periostracum by papillae and protoplasmicstrands which persist during mineral deposition and ensure thatboth shell and attached mantle remain in situ relative to theoutwardly expanding inner surface of the outer mantle lobe.In the prototypic brachiopod, the lobate cells are likely atfirst to have occupied the hinge of the mantel fold but laterto have been displaced into their present position by the rigidoutward growing edge of the mineral shell.     

Cyclic variations of stomatal aperture observed under the scanning electron microscope

Scanning electron micrographs taken from a vertical positionrevealed the stomatal aperture at the central pore and at theouter ledge. Cyclic variations in leaf temperature of tobacco (Nicotianatabacum cv. MC) and sunflower (Helianthus annuus cv. Large Russian)after illumination coincided with the aperture at the centralpore, but did not always at the outer ledge. The relative turgidityof leaf was inversely proportional to the aperture at the outerledge. Moreover, the opening at the central pore was an "8"-shapedaperture provided the opening was small. Thus, the present experimentsindicated the importance in measurement of the stomatal apertureat the central pore for quantitative and qualitative determinationof cyclic variations in the stomatal aperture. (Received October 7, 1977; )     

Crystalline structure,orientation and nucleation of the nacreous tablets in the cephalopod <Emphasis Type="Italic">Nautilus</Emphasis>

The nacreous tablets in the Nautilus shell have similar crystalline structure as the tablets in the gastropod Gibbula shell. Etching with Mutvei’s solution reveals that each tablet is composed of vertical crystalline columns that are structurally similar to the acicular crystallites in the outer spherulitic-prismatic layer of the shell wall. The columns are attached to each other to form numerous vertical crystalline lamellae, oriented parallel to the longitudinal axis of the tablet. It is still unknown whether or not the orientation of the vertical lamellae corresponds to that of the crystallographic a- or b-axis. The orientation of the crystalline lamellae in the adjacent tablets is parallel in some nacreous laminae, but random in other laminae. Similar large variation was found in the nacreous tablets of the gastropod and bivalve shells. The nucleation sites of the nacreous tablets are predominantly situated on the peripheral portion of the upper surface of the preceding tablet, both in the shell wall and septa. The “aragonite-nucleating proteins” in the central portion of the crystal imprints of the organic interlamellar sheets, described by several writers, have therefore a negative correlation with the nucleation sites of the nacreous tablets.     

Oxygen and CO2 Exchange and Acid-Base Regulation in the Avian Embryo

The avian embryo exchanges the oxygen and carbon dioxide withthe ambient air by diffusion. The respiratory organ is the chorioallantois,endowed with a rich circulation. Between ambient air and chorioallantoiccapillary blood are interposed the porous shell fibrous shellmembranes, and the chorioendothelium which compose the diffusionbarrier. The air cell is formed between the two shell membranesin the blunt end of the egg. The diffusion barrier is dividedinto an outer barrier (shell plus outer membrane) and an innerbarrier (inner membrane plus chorioendothelium and capillaryblood). The resistance to gas diffusion (the reciprocal of thediffusive conductance) in the outer barrier is almost fixedthroughout incubation while that in the inner barrier decreasesas the embryo develops. Because of the fixed outer barrier conductance,the embryo is obliged to take up oxygen under hypoxic conditionsagainst increasing metabolism with development and encountersa relative respiratory acidosis. In connection with the diffusivehypoventilation caused by the fixed outer barrier conductancethe respiratory factors of the allantoic circulation changeprogressively with development to moderate the restraint ofgas exchange through the shell. Blood oxygen capacity and hemoglobinincrease with development in association with an increase inerythrocyte count and hematocrit value. In addition, a progressiveleftward shift of the oxygen dissociation curve occurs. Theincreases in the allantoic blood flow and chorioallantoic capillaryvolume contribute to the increasing conductance of the innerbarrier. Furthermore regulation of acid base balance is inferredin the developing embryo.     

The phosphate-rich cover on the eggshells of grebes (Aves: Podicipitiformes)

Spheres of a nut-like morphology–a "kernel" formed from spherules contained in a "shell" of globular sub-units–were a unique feature of the cover on the eggshells of six species of the order Podicipitiformes: Red-necked grebe (Podiceps grisegend) , Great crested grebe (Podiceps cristatus) , Little grebe (Tachybaptus ruficollis) , Black-necked grebe (Podiceps nigrkollis) , Slavonian grebe (Podiceps auritus) and Pied-billed grebe (Podilymbus podiceps). The spheres did not give a pattern with X-ray diffraction; the innermost ones were embedded in the dense crystalline layer on the outer surface of the calcitic portion of the shell. Electron-probe microanalysis revealed that the cover of spheres contained principally oxygen, calcium, phosphorus and sulphur and infra-red analysis demonstrated the presence of phosphate. The outer surface of the stratum of spheres was bounded by a thin, fissured layer of amorphous material.
These observations raise questions about the ionic environment obtaining in the shell gland towards the end of shell formation and the adaptive significance of the cover on the eggshells of members of the Podicipitiformes.     

Gas Exchange of the Avian Egg Time, Structure, and Function

Data are presented for oxygen consumption water loss duringincubation water vapor conductance of the shell and pore numberof avian eggs and the way in which these values relate not onlyto egg mass but also to incubation time. It is proposed thatall these functions are proportional to the product of egg massand rate of development where the latter is defined as the inverseof incubation time. These interrelationships account at theend of incubation for similar O₂ and CO₂ tensions in the airspace of eggs utilization of calories (0.5 kcal g^–1) andwater loss (15 g g^–1)     

Fine Structure of the Contractile Vacuole Pore in Paramecium*

The pore through which a Paramecium contractile vacuole communicates with the external environment is a 1.2 μm long and 1 μm diameter cylindrical orifice in the pellicle. During diastole, the vacuole:pore junction is closed by a substantial diaphragm which parts to the side at systole. The diaphragm is composed of inner and outer membranes continuous with the vacuole and pore membranes, respectively, and an intervening cytoplasmic layer containing filaments and irregular membranous tubules and vesicles. Microtubules, organized into 2 sets, are an important component of the pore apparatus. One set of ～ 16 microtubules forms an annulus around the pore. These microtubules are organized into a right-handed helix with a pitch of 0.5-0.6 μm, and thus complete slightly more than 2 turns in their course from the level of the diaphragm to the pore outer lip. They appear to be embedded in a layer of dense material immediately adjacent to the pore membrane. The other set consists of 5 or more bands of 10–20 microtubules which radiate in a slight left-handed helix from an insertion at the pore out over the vacuole surface to the ampullae.     

Studies on the egg shell (oviducal and oviposited) of Chelonia mydas L.

The outer calcified surface of the turtle egg shell consists primarily of crystalline aggregates of calcium carbonate in its aragonite form, together with a small amount (< 5 %) of calcitic material. The latter is first deposited to be followed by aragonite deposition.In the first instance, calcification occurs on the rims of discrete pits formed by the lateral deflection of the ends of soft shell membrane fibres. As crystal deposition continues these pits become filled in and eventually occluded.Micro- and X-ray diffraction analyses of the calcified layer indicate the presence of phosphorus and sulphur. The effects of these elements on the type of crystal deposited, (i.e., aragonite or calcite) is discussed.     

Resource assessment in hermit crabs: the worth of their own shell

Animals gather information about the quality of a resource throughits assessment and behave accordingly as a result of adaptivemotivational changes. In the hermit crab Pagurus longicarpus,we investigated whether an individual was affected in its motivationto acquire a new shell by the quality of the domicile shell(own resource value [ORV]), of the offered shell (external resourcevalue [ERV]), or of both and asked whether its motivation wasaltered by the information gathered during shell investigation.We analyzed the behavior of hermit crabs inhabiting shells ofdiffering qualities and compared their willingness to acquirean offered shell—optimal, smaller than optimal, or largerthan optimal—by measuring the latency to approach it,the number of shell investigation, and its total duration. Crabsin smaller shells (SSs) approached more quick and often theoffered shell, whereas crabs in larger shells investigated theoffered shell more thoroughly. The readiness of crabs to approachthe offered shell and the extent of its investigation were independentof the ERV but were exclusively affected by the ORV, whereasthe number and duration of shell investigation did not changewith time as investigation proceeded, except for crabs in SSs.These results suggest that P. longicarpus' motivation to acquirea new shell is exclusively influenced by the value of the shellit inhabits rather than by the quality of the shell it is offeredand that this species does not gather—or does not use—informationabout ERV during investigation.     

Dynamics of dolphin porpoising revisited

Porpoising is the popular name for the high-speed surface piercingmotion of dolphins and other species, in which long, ballisticjumps are alternated with sections of swimming close to thesurface. The first analysis of this behavior (Au and Weihs,1980) showed that above a certain "crossover" speed this behavioris energetically advantageous, as the reduction in drag dueto movement in the air becomes greater than the added cost ofleaping. Since that publication several studies documented porpoisingbehavior at high speeds. The observations indicated that thebehavior was more complex than previously assumed. The leapswere interspersed with relatively long swimming bouts, of abouttwice the leap length. In the present paper, the possibilityof dolphins using a combination of leaping and burst and coastswimming is examined. A three-phase model is proposed, in whichthe dolphin leaps out of the water at a speed U_f, which is thefinal speed obtained at the end of the burst phase of burstand coast swimming. The leap is at constant speed and so theanimal returns to the water at U_f, goes to a shallow depth andstarts horizontal coasting while losing speed, till it reachesU_i. At that point it starts active swimming, accelerating toU_f. It then starts the next leap. Ranges of speeds for whichthis three-stage swimming is advantageous are calculated asa function of animal and physical parameters. Notation C—Constant defined in equation (12) C_D—Coasting drag coefficient D—Drag g—Gravitational acceleration H—Height of jump J—Energy required for jump k—Ratio of swim length to jump length l—Distance L—Total distance (eq. 28) m—Added mass M—Animal mass M₁—Total mass r—Coefficient defined in eq. (22) R—Ratio of energies, for three-phase swimming R₂—Ratio of energies, for burst and coast swimming t—Time T—Thrust U—Speed V—Body volume W—Weight     

REMOTE BIOMINERALIZATION IN DIVARICATE RIBS OF STRIGILLA AND SOLECURTUS (TELLINOIDEA: BIVALVIA)

Deposits composed of aragonite prisms, which were formed afterthe outer shell layer, have been found at the posterior steepslopes of divaricate ribs in two species of Strigilla and anothertwo of Solecurtus. These prisms have their axes oriented perpendicularto the outer shell surface and differ in morphology from fibresof the surface-parallel composite prisms forming the outer shell.They display crystalline features indicating that, unlike crystalsforming the outer shell surface, their growth front was free,unconstrained by the mantle or periostracum. These particulardeposits are called free-growing prisms (FGPs). In these generathe periostracum is clearly not the substrate for biomineralizationand, upon formation, does not adhere to the steep slope of ribs,but detaches at the rib peak and reattaches towards the posterior,just beyond the foot of the posterior scarps of ribs. In thisway, a sinus or open space developed between the internal surfaceof the periostracum and the outer shell surface along each steeprib slope. These spaces could remain filled with extrapallialfluid after the mantle advances beyond that point during shellsecretion. FGPs grow within this microenvironment, out of contactwith the mantle. Other species with divaricate ribs do not developFGPs simply because the periostracum adheres tightly to both ribslopes (which are never so steep as in Solecurtus and Strigilla).FGPs constitute one of the rare cases of remote biomineralizationin which aragonite is produced and direct contact with the mantlenever takes place. (Received 22 November 1999; accepted 20 February 2000)     

隆线溞[Daphnia(Ctenodaphnia)carinata]卵鞍的超微结构

在不利的环境条件下,枝角类中有一部分种类可以形成卵鞍(ephippium),内含休眠卵。本文应用扫描电镜和透射电镜对隆线溞的卵鞍进行了超微结构的研究。研究表明:卵鞍外面大部分略呈浅的蜂窝状,内面则排布着多数卵石状小突起。卵鞍分为内外两层,两层的超微结构截然不同;各层又可分为三小层。     

Structure of shells of eggs of Callisaurus draconoides (Reptilia, Squamata, Iguanidae)

Female zebra-tailed lizards (Iguanidae: Callisaurus draconoides ) lay roughly ovoid eggs with thin, highly extensible shells. The outer surface of the eggshell is a thin, calcareous crust of calcium carbonate in the calcite morph. Immediately beneath the crystalline matrix is a shell membrane composed of multiple layers of fibres organized into an undulating series of troughs and crests, apparent in both cross-section and surface view. The outer surface of the shell membrane is differentiated into a tightly woven fibrous mat that may serve to anchor the calcareous layer to the membrane. Organization of the eggshell into a series of troughs and crests serves to increase the surface area available for contact with the substrate and, presumably, to increase the capacity of the eggshell to stretch as the egg absorbs water.     

Separation and Properties of the Inner and Outer Membranes of a Psychrophilic Bacterium, Vibrio sp. Strain ABE-1

The inner and outer membranes from a marine psychrophilic bacterium,Vibrio sp. ABE-1, were separated and purified by desalting,lysozyme treatment, and ultracentrifugation. Enzyme assay, electrophoresis,and chemical analysis showed that they had been highly purified.The isolated outer membrane had no 3-deoxy-D-mannooctulosonate(KDO) and chemical analysis revealed that it contained aboutthree times as much total lipid as the inner membrane. Consequently,the density of the inner membrane was higher than that of theouter membrane (1.19–1.23 gµcm³ for IM; 1.16–1.20g–cm³ for OM). The absolute majority of the lipids inthese membranes were phospholipids, more than half of whichwas identified as phosphatidylethanolamine. Virtually all cardiolipinwas found exclusively in the inner membrane, whereas an unidentifiedphospholipid containing hexosamine was only in the outer membrane.The ratios of unsaturated to total fatty acids in the innerand outer membranes were 76.0% and 69.6%, respectively. (Received February 23, 1984; Accepted August 8, 1984)     

Larval experience and latent effects--metamorphosis is not a new beginning

For many years ecologists have documented the remarkable within-speciesvariation inherent in natural systems—for example, variabilityin juvenile growth rates, mortality rates, fecundities, timeto reproductive maturity, the outcomes of competitive interactions,and tolerance to pollutants. Over the past 20 years, it hasbecome increasingly apparent that at least some of this variationmay reflect differences in embryonic or larval experiences.Such experiences may include delayed metamorphosis, short termstarvation, short term salinity stress, or exposure to sublethalconcentrations of pollutants or sublethal levels of ultra violetirradiation. Latent effects—effects that have their originsin early development but that are first exhibited in juvenilesor adults—have now been documented among gastropods, bivalves,echinoderms, polychaetes, crustaceans, bryozoans, urochordates,and vertebrates. The extent to which latent effects alter ecologicaloutcomes in natural populations in the field, and the mechanismsthrough which they are mediated are largely unexplored.