Studies in the biochemistry of cirripede eggs. VI. Changes in the general biochemical composition during development of Tetraclita squamosa rufotincta Pilsbry,Balanus perforatus Brug., and Pollicipes cornucopia Darwin

The carbohydrate, protein, lipid, total RNA and DNA at four stages in the development of the eggs of Tetraclita squamosa rufotincta Pilsbry, Balanus perforatus Brug., and Pollicipes cornucopia Darwin have been determined. The initial caloric content varies with species, that of the large egg of Tetraclita being extremely high; on a relative volume-specific basis, however, differences, although still present, are much less variable. A comparison has been made with other species. Those eggs in which lipid predominates over protein in Stage I lose more lipid during development. The relatively large quantities of reserves in Tetraclita may serve as metabolic substrate during the development of the planktonic larvae under nutrient-poor conditions.     

Scanning Electron Microscopy of Mouth Appendages in Six Species of Barnacles (Crustacea Cirripedia Thoracica)

The morphology and setation of mouth appendages (trophi) are investigated with scanning electron microscopy in the balanomorph barnacles Semibalanus balanoides, Balanus balanusand B. nubilusand in the pedunculate barnacles Pollicipes polymerus, P. cornucopiaand Lepas anatifera.It is difficult to uphold a clearcut distinction between denticles and setae and several types of setae also intergrade with each other. The trophi of Pollicipes polymerusand P. cornucopiahave the most ‘generalized’ morphology. The palps of Pollicipesand L. anatiferaare simpler than those in balanomorphs and carry a single type of seta. In L. anatiferathe mandibles and maxillules are adapted to a semi-predatory feeding habit by carrying large, pointed teeth, but this species lacks the foliate-serrate setae which populate the palps in the other species studied. Compared with Pollicipesand Lepas, the three balanomorphs have palps with a complex setation. The trophi in S. balanoidesdiffer from the two species of Balanusin numerous features of setation and denticulation, notably in having the palps populated by plumodenticulate setae homologous to purely denticulated types in Balanus.It is suggested that SEM studies of barnacle trophi will provide characters of use in estimating phylogeny.     

Costs and benefits of maternal behaviour in the green lynx spider (Oxyopidae,Peucetia viridans)

I examined the adaptiveness of maternal behaviour in the green lynx spider Peucetia viridans (Hentz), by measuring the costs and benefits to the female of egg-sac tending. P. viridans guards her egg sac for 6–8 weeks until the young have emerged and dispersed. I removed females from egg sacs in the field either immediately after oviposition or after they had guarded the egg sacs for 2 weeks, and compared the fate of these sacs with that of control sacs which had females present throughout. Survival rates of egg sacs were significantly improved by the presence of a female. The major sources of mortality were ant predation and sac dislodgement, both of which were significantly reduced by a guarding female, and mantispid brood parasitism, which was not affected by the female. To measure the spiders' lifetime reproductive output, I allowed caged females to construct egg sacs. Half had their first egg sac removed shortly after oviposition, while half were allowed to guard. Significantly more non-guarding than guarding females produced second egg sacs. Thus for the green lynx spider, egg sac guarding decreases lifetime egg production, but increases offspring survival. Under north Florida conditions, the benefits of guarding outweight the costs, and the females have a higher lifetime reproductive success than if they did not guard.     

Egg sac damage and previous egg sac production influence truncated parental investment in the wolf spider,Pardosa milvina

Life history theory predicts that iteroparous animals adaptively partition reproductive effort between current and future reproduction. When rearing costs of current offspring exceed the potential benefits, parental care should be terminated and deferred toward future reproduction. We tested two related predictions that follow from life history theory: (a) parents should be sensitive to offspring viability and withhold parental care if offspring survival probability drops and future reproductive opportunities are likely, and (b) parents should be less sensitive to offspring survival probability when future reproduction is unlikely and maximize parental care late in life. The wolf spider, Pardosa milvina, demonstrates extensive parental care; however, they may also abandon or cannibalize their egg sacs. We tested the effects of egg sac damage and production of a previous egg sac on egg sac abandonment and cannibalism decisions. Among four egg sac groups (1st egg sac intact, 1st egg sac damaged, 2nd egg sac intact, 2nd egg sac damaged), we daily monitored egg sac abandonment and cannibalism and measured differences in egg sac searching, protection, and grooming among removed and damaged egg sacs (N = 116 with 1st egg sac and 88 with 2nd egg sac). Females with first egg sacs abandoned and cannibalized damaged egg sacs significantly more compared to unmanipulated egg sacs; however, females with second egg sacs were insensitive to egg sac damage. Females also spent significantly more time protecting second egg sacs compared to first egg sacs and groomed damaged egg sacs significantly more than undamaged. These results support the general predictions of life history theory that indicate that abandonment and cannibalism should decrease with diminished future reproductive potential and that parents should be less sensitive to indicators of offspring survival probability late in life.     

Egg-laying of the golden silk spider, Nephila clavipes L. (Araneae,Araneidae): Functional analysis of the egg sac

The egg-laying sequence of the orb-weaving spider Nephila clavipes was observed in the field setting. The functions of certain modal characteristics of the egg sac were assessed through the collection of normative field data on overwintering egg sacs and through the effects of egg sac manipulations on spiderling production. Components of the egg sac were manipulated before and after egg hatching to determine if they promoted offspring survival at different times in development. Naturally occurring and induced changes in the position and structure of the egg sac resulted in similar decreases in productivity. Certain components of the egg sac were important primarily before hatching, and others throughout the entire overwintering period. These results and the limitations of functional analyses of behaviours are discussed.     

Aspects of the ecology and energetics of the egg sac parasites of the wolf spider Pardosa lugubris (Walckenaer)

Summary The egg sacs of the wolf spider Pardosa lugubris are parasitised by two ichneumonid wasps, namely Gelis micrurus and Hidryta sordidus. The parasites are bivoltine, the first generation parasitising the summer egg sac of P. lugubris and the second generation parasitising, and overwintering in, the autumn egg sac. Parasitised egg sacs have a characteristic appearance.The degree of egg sac parasitism was 9.8% and 6.9% for the two egg sac batches in 1965 and 2.9% and 34.8% for those of 1966. The degree of parasitism varies from area to area as does the percentage parasitism attributable to the two species.The gross growth efficiency for Hidryta from the first egg sac is 31.2% and from the second 16.2%. 3.0% in 1965 and 4.3% in 1966, of the annual yield from P. lugubris to predators and parasites, goes to the egg sac parasites. This represents reductions in recruitment of 9.1% in 1965 and 10.9% in 1966.     

Maternal defensive behaviors of <Emphasis Type="Italic">Uloborus</Emphasis> sp. (Araneae,Uloboridae): behavioral repertoire and influence of clutch size and female size on female aggressiveness

Maternal care in spiders often involves behaviors associated with the protection of eggs and spiderlings against parasitoids and predators (including conspecifics). The females of several species have been documented to move their egg sacs away from natural enemies or to invest in active defense behaviors against web invaders, such as parasitoid wasps or araneophagic spider species, to protect their brood. In this study, we present observations of protective behavior by Uloborus sp. females carrying egg sacs. We also investigated whether brood size and female size influence female aggressive behaviors and response time against an artificial source of disturbance. Females carrying egg sacs almost immediately perceived and reacted aggressively against the artificial stimulus, whereas females without egg sacs moved away or ran to the web margins, avoiding the source of disturbance. The aggressive response was independent of clutch size and female body size, indicating that all females will risk interacting with potential agents of egg mortality. This systematic response by all females with egg sacs may be important for reducing the incidence of attack by the egg predator wasp Bathyzonus sp. (Ichneumonidae).     

Studies on the reproduction of cirripedes. II. Setting of the lamellae; action of protease and disintegration of the oviducal sac

The possible modes of breakdown of the oviducal sac which comes to surround the egg lamellae in cirripedes, has been investigated in a variety of common species. Bacterial action appears to be unimportant.The effect of protease on the sac has been investigated. The rate of breakdown is unrelated to season when it is considered relative to the stage of embryonic development. Apart from Verruca stroemia Müller which is not attacked, the initial rate of ‘dissolution’ is rapid, reaches a minimum, and then increases. Although the time course is similar in all the species examined, there are quantitative differences between species.The differences may only reflect differences in the protein composition of the sac and their resistance to protease. Under the action of its own enzyme species may show much less differences. It is considered that the results indicate that the enzyme necessary for proteolysis is produced by the embryos themselves at a certain stage in their development. The results are discussed relative to other known hatching enzymes.     

Relative investment in egg load and poison sac in fig wasps: Implications for physiological mechanisms underlying seed and wasp production in figs

Fig pollinating wasps and most non-pollinator wasps apply secretions from their poison sacs into oviposited flowers that appear necessary to the formation of the galls that their developing offspring consume. Thus, both eggs and poison sac secretions appear to be essential for wasp reproduction, but the relative investment in each is unknown. We measured relative investment in poison sac and egg production in pollinating and non-pollinating wasps associated with seven species of monoecious Panamanian figs representing both active and passive pollination syndromes. We then collected similar data for four fig hosts in China, where some wasp species in the genus Eupristina have lost the ability to pollinate (“cheaters”). All wasps examined possessed large poison sacs, and we found a strong positive correlation between poison sac size and absolute egg production. In the Panamanian species, the relative poison sac to egg investment was highest in the externally ovipositing non-pollinator wasps, followed by active pollinators, then by passive pollinators. Further, pollinator wasps of fig species with demonstrated host sanctions against “cheating” wasps showed higher investment in the poison sac than wasps of species without sanctions. In the Chinese samples, relative investment in the poison sac was indistinguishable between pollinators and “cheaters” associated with the same fig species. We suggest that higher relative investment in poison sac across fig wasp species reflects higher relative difficulty in initiating formation of galls and subsequently obtaining resources from the fig. We discuss the implications for the stability of the fig–wasp mutualism, and for the ability of non-pollinators to exploit this mutualism.     

Behaviour of colonial and solitary spiders of the theridiid species Anelosimus eximius

Most Anelosimus eximius live in colonies, but a few females emigrate short distances and establish small, individual webs. Field studies were conducted on one colony and nearby smaller webs in order to describe communal activities and division of labour, and to note costs and benefits of remaining in the parental colony and emigrating. Adult and juvenile females repaired the web and captured prey. Adult females, rather than juveniles, cared for egg sacs and fed young. When colonial egg sacs were abundant, females moved from sac to sac. Several colonial females regurgitated food to spiderlings which had recently emerged from a particular sac. In smaller webs inhabited by two or three females, the mother cared for the sac but all females fed the young by regurgitation. Males rarely participated in communal activities. Advantages of colony living include protection from predators, the availability of large prey and, in the event of a female's death, the care of her egg sac and feeding of her young. Cannibalism of the egg sac is a potential cost of coloniality, affecting about 10–20% of colonial sacs. Cannibalism was not observed in the smaller webs. However, the costs of emigrating are very high: most of the solitary females disappeared, leading to interspecific predation on their eggs and young.     

Assessing the In Situ Fertilization Status of Two Marine Copepod Species,Temora longicornis and Eurytemora herdmani; How Common Are Unfertilized Eggs in Nature?

We utilized an egg staining technique to measure the in situ fertilization success of two marine copepod species, Temora longicornis and Eurytemora herdmani from May to October 2008 in coastal Maine and correlated fertilization success with environmental conditions in their habitat. T. longicornis is a free spawning species that releases eggs into the ambient seawater after mating. In contrast, E. herdmani carries eggs in an egg sac until they hatch. The proportion of fertilized eggs within E. herdmani egg sacs was significantly higher than the freely spawned clutches of T. longicornis. This may be a result of the asymmetrical costs associated with carrying vs. spawning unfertilized eggs. T. longicornis frequently laid both fertilized and unfertilized eggs within their clutch. T. longicornis fertilization was negatively associated with chlorophyll concentration and positively associated with population density in their local habitat. The fertilization status of E. herdmani egg sacs was high throughout the season, but the proportion of ovigerous females was negatively associated with an interaction between predators and the proportion of females in the population. This study emphasizes that, in addition to population level processes, community and ecosystem level processes strongly influence the fertilization success and subsequent productivity of copepods.     

An arboreal spider protects its offspring by diving into the water of tank bromeliadsUne araignée arboricole utilise les réservoirs d’eau des broméliacées pour se protéger et protéger son cocon

Cupiennius salei (Ctenidae) individuals frequently live in association with tank bromeliads, including Aechmea bracteata, in Quintana Roo (Mexico). Whereas C. salei females without egg sacs hunt over their entire host plant, females carrying egg sacs settle above the A. bracteata reservoirs they have partially sealed with silk. There they avoid predators that use sight to detect their prey, as is known for many bird species. Furthermore, if a danger is more acute, these females dive with their egg sacs into the bromeliad reservoir. An experiment showed that this is not the case for males or females without egg sacs. In addition to the likely abundance of prey found therein, the potential of diving into the tank to protect offspring may explain the close association of this spider with bromeliads. These results show that, although arboreal, C. salei evolved a protective behavior using the water of tank bromeliads to protect offspring.     

Comparative analysis of stalked and acorn barnacle adhesive proteomes

Barnacles interest the scientific community for multiple reasons: their unique evolutionary trajectory, vast diversity and economic impact—as a harvested food source and also as one of the most prolific macroscopic hard biofouling organisms. A common, yet novel, trait among barnacles is adhesion, which has enabled a sessile adult existence and global colonization of the oceans. Barnacle adhesive is primarily composed of proteins, but knowledge of how the adhesive proteome varies across the tree of life is unknown due to a lack of genomic information. Here, we supplement previous mass spectrometry analyses of barnacle adhesive with recently sequenced genomes to compare the adhesive proteomes of Pollicipes pollicipes (Pedunculata) and Amphibalanus amphitrite (Sessilia). Although both species contain the same broad protein categories, we detail differences that exist between these species. The barnacle-unique cement proteins show the greatest difference between species, although these differences are diminished when amino acid composition and glycosylation potential are considered. By performing an in-depth comparison of the adhesive proteomes of these distantly related barnacle species, we show their similarities and provide a roadmap for future studies examining sequence-specific differences to identify the proteins responsible for functional differences across the barnacle tree of life.     

Studies in the biochemistry of cirripede eggs. IV. The free Amino-acid pool in the eggs of Balanus balanoides (L.) and B. balanus (L.) during development

Changes in the ‘free’ amino acids, betaine, and trimethylamine oxide during the development of the eggs of Balanus balanoides (L.) and B. balanus (L.) have been determined; the results are given in terms of μM/g dry wt, μM/g water, and μM/10⁶ eggs. The amino acids are derived from the yolk proteins the net composition of which is known. Free amino acids are present in considerable quantity, as is commonly the case with crustacean tissue. Changes in the individual amino acids are discussed. B. balanus eggs contain large, and relatively constant, amounts of sarcosine; its function is unknown but large quantities are present in the more highly evolved cirripedes so far examined. A possible relation between betaine glycine, and sarcosine relative to choline metabolism is considered. Large amounts of taurine are present. There is a striking increase in β-alanine in the late stages of development; in B. balanoides it comes to be the most, and in B. balanus the third most common amino acid; its possible involvement in purine metabolism is considered. The relation between the amounts of the various entities in the eggs and in the bodies of the adult are examined.     

Studies in the biochemistry of cirripede eggs. V. Changes in the general biochemical composition during development of Chthamalus stellatus (Poli)

The biochemical composition of the eggs of Chthamulus stellatus (Poli) during their development has been investigated. The eggs are incubated within the mantle cavity of the adult but no substrates are contributed by the latter. Carbohydrates, protein, and lipid are all utilized. In contrast to two cold-water species, Balanus balanoides and B. balanus perviously investigated lipid rather than protein is consumed during the early stages. Possible explanations for this are considered.     

Factors influencing fighting and threat in the parrot genus Trichoglossus

Parrots of the genus Trichoglossus perform elaborate repertoires of visual threat displays during agonistic interactions. The displays of different species and subspecies differ in complexity and composition. As repertoire complexity increases, taxa tend to emphasize the more exaggerated components of display, and they also show a reduced inclination to attack a mirror-image opponent. These covarying attributes of threat displays in Trichoglossus are highly correlated with interspecific differences in bill-length. These findings are discussed in the light of recent theoretical advances in the study of animal threat and fighting behaviour.     

The vocal sac of Hylodidae (Amphibia,Anura): Phylogenetic and functional implications of a unique morphology

Anuran vocal sacs are elastic chambers that recycle exhaled air during vocalizations and are present in males of most species of frogs. Most knowledge of the diversity of vocal sacs relates to external morphology; detailed information on internal anatomy is available for few groups of frogs. Frogs of the family Hylodidae, which is endemic to the Atlantic Forest of Brazil and adjacent Argentina and Paraguay, have three patterns of vocal sac morphology—that is, single, subgular; paired, lateral; and absent. The submandibular musculature and structure of the vocal sac mucosa (the internal wall of the vocal sac) of exemplar species of this family and relatives were studied. In contrast to previous accounts, we found that all species of Crossodactylus and Hylodes possess paired, lateral vocal sacs, with the internal mucosa of each sac being separate from the contralateral one. Unlike all other frogs for which data are available, the mucosa of the vocal sacs in these genera is not supported externally by the mm. intermandibularis and interhyoideus . Rather, the vocal sac mucosa projects through the musculature and is free in the submandibular lymphatic sac. The presence of paired, lateral vocal sacs, the internal separation of the sac mucosae, and their projection through the m. interhyoideus are synapomorphies of the family. Furthermore, the specific configuration of the m. interhyoideus allows asymmetric inflation of paired vocal sacs, a feature only reported in species of these diurnal, stream‐dwelling frogs.     

Spiderling sex ratio and maternal investment in the bolas spider Mastophora cornigera (Araneae,Araneidae)

Male Mastophora cornigera exit egg sacs as adults, which allowed us to determine spiderling sex ratios and patterns of maternal investment in this species. We collected 15 egg sacs produced by seven mothers, which yielded 1945 emergent spiderlings which were sexed, 1850 of which were weighed. Two emergent broods were significantly male and female biased and were unaffected by pre-emergence mortality. The weights of male and female spiderlings differed in eight broods, with males and females being heavier in four cases each. Five of these broods were derived from multiple egg sac sets produced by one mother, and in each case, the total mean male and female spiderling weights for all broods in a set were biased in the same direction as the biased brood(s) within that set. Mean emergent spiderling weight was independent of brood size and sex ratio for both males and females. Despite such independence, sex allocation in M. cornigera can favor sons, daughters, or both equally, and by numbers, by weight, or both at once. The proximate mechanisms and adaptive significance of such variability is unknown. We also review evidence for gender-biased allocations in arachnid offspring and suggested mechanisms for their applicability to M. cornigera.     

Isolation and preservation of the living embryo sac ofCrinum asiaticum L. var.japonicum baker

The enzymatic maceration method was used to isolate an intact embryo sac ofCrinum asiaticum and its component cells. Best results were obtained when using enzyme solutions that contained pectinase hemicellulase, cellulase and pectolyase. Aseptic ovules were incubated in the enzyme solution for 1.5 hr at 25 C. This allowed the isolation of embryo sacs to yield up to 20% of the amount present. An isolated embryo sac usually consists of an egg cell, synergids, antipodals and a central cell. Some embryo sacs can be digested as gametophytic protoplast. The size, shape and position of the isolated embryo sac seemingly possessed similarities with those of the fixed embryo sac in the ovary. An isolated embryo sac can be in a living state when the result of the fluorochromatic reaction (FCR) and protoplasmic streaming is positive. When cultured in proper media, 68% of the isolated gametophytic protoplasts were observed to have sustained their positive FCR for more than 1 month.     

Lysozyme in egg whites of tortoises and turtle. Purification and properties of egg white lysozyme of Trionyx gangeticus Cuvier.

Egg white proteins of three species of tortoises and turtle and of hen have been compared by electrophoretic and immunochemical methods. The proteins lacked similarity in electrophoresis, but tortoise and turtle egg white proteins which did not crossreact with those of the hen showed some cross-reaction among themselves. The occurrence of lysozyme as two allelic variants which were distinguishable in electrophoresis was noted only in the egg white of one of the species of tortoise, namely, Trionyx gangeticus Cuvier. Tortoise lysozyme which showed strong lytic activity toward cell walls of Micrococcus lysodeikticus did not exhibit any cross-reaction with hen lysoyzme. It was purified, crystallized, and found to be homogeneous in sodium dodecyl sulfatepolyacrylamide gel electrophoresis, immunochemical tests, and sedimentation. The physicochemical and enzymatic properties of tortoise lysozyme were found to be strikingly similar to those of hen lysozyme with minor differences which could be due to differences in their primary structure. Its average molecular weight of 15,400 was determined from sedimentation and diffusion coefficient values, Archibald experiment, and amino acid composition. The molecule appeared to undergo pH-dependent expansion at pH 2 and dimerization above pH 5.7. In enzymatic properties, tortoise lysozyme showed a specific activity of 29,000–31,000 units and gave a pH optimum at pH 7.5 and an apparent K_a value of 250 mg· liter^?1. Like hen lysozyme, its activity showed strong ionic strength dependence, weak chitinase activity, susceptibility to inhibition by N-acetyl-glucosamine, and stability toward heat.