Physical features and chitin content of eggs from the mosquito vectors Aedes aegypti,Anopheles aquasalis and Culex quinquefasciatus: Connection with distinct levels of resistance to desiccation

Mosquito eggs are laid in water but freshly laid eggs are susceptible to dehydration, if their surroundings dry out at the first hours of development. During embryogenesis of different mosquito vectors the serosal cuticle, an extracellular matrix, is produced; it wraps the whole embryo and becomes part of the eggshell. This cuticle is an essential component of the egg resistance to desiccation (ERD). However, ERD is variable among species, sustaining egg viability for different periods of time. While Aedes aegypti eggs can survive for months in a dry environment (high ERD), those of Anopheles aquasalis and Culex quinquefasciatus in the same condition last, respectively, for one day (medium ERD) or a few hours (low ERD). Resistance to desiccation is determined by the rate of water loss, dehydration tolerance and total amount of water of a given organism. The ERD variability observed among mosquitoes probably derives from diverse traits. We quantified several attributes of whole eggs, potentially correlated with the rate of water loss: length, width, area, volume, area/volume ratio and weight. In addition, some eggshell aspects were also evaluated, such as absolute and relative weight, weight/area relationship (herein called surface density) and chitin content. Presence of chitin specifically in the serosal cuticle as well as aspects of endochorion external surface were also investigated. Three features could be related to differences on ERD levels: chitin content, directly related to ERD, the increase in the egg volume during embryogenesis and the eggshell surface density, which were both inversely related to ERD. Although data suggest that the amount of chitin in the eggshell is relevant for egg impermeability, the participation of other yet unidentified eggshell attributes must be considered in order to account for the differences in the ERD levels observed among Ae. aegypti, An. aquasalis and Cx. quinquefasciatus.     

The cooling time of fertile chicken eggs at different stages of incubation

We asked whether or not the thermal characteristics of fertile avian eggs changed throughout incubation. The cooling and warming times, expressed by the time constant τ of the egg temperature response to a rapid change in ambient temperature, were measured in fertile chicken eggs at early (E7), intermediate (E11) and late (E20) stages of embryonic development. Same measurements were conducted on eggs emptied of their content and refilled with water by various amounts. The results indicated that (1) the τ of a freshly laid egg was ~50 min; (2) τ decreased linearly with the drop in egg water volume; (3) the dry eggshell had almost no thermal resistance but its wet inner membrane contributed about one-third to the stability of egg temperature; (4) the egg constituents (yolk, albumen and embryonic tissues) and the chorioallantoic circulation had no measurable effect on τ; (5) the presence of an air pocket equivalent in volume to the air cell of fertile eggs reduced τ by about 3 min (E7), 5 min (E11) and 11 min (E20). Hence, in response to warming the egg τ at E20 was slightly shorter than at E7. In response to cooling, the egg τ at E20 was similar to, or longer than, E7 because embryonic thermogenesis (evaluated by measurements of oxygen consumption during cold) offset the reduction in τ introduced by the air cell. In conclusion, until the onset of thermogenesis the thermal behavior of a fertile egg is closely approximated by that of a water-filled egg with an air volume equivalent to the air cell. It is possible to estimate the cooling τ of avian eggs of different species from their weight and incubation time.     

Organic,inorganic, and caloric composition of eggs,pentaculae, and adults of the brooding sea cucumber Cucumaria curata cowles (Echinodermata: Holothuroidea)

Levels (percentage composition) of water, ash, carbohydrate, lipid, protein, and calories were determined for eggs, pentaculae, and adults of the sea cucumber Cucumaria curata Cowles. Component contents (μg/individual) were calculated for eggs and pentaculae. During the 28 days of development to hatching, the large yolky eggs gain water and ash, the total dry weight increasing from 169 to 190 μg/egg during embryogenesis. There were no statistically significant changes in lipid, protein, and caloric contents during embryogenesis, but carbohydrate decreased by 0.82 μg/egg.The decrease in carbohydrate is sufficient to account for estimated embryonic energy requirements. Based on the utilization of carbohydrate, embryos of C. curata show a nutritional pattern similar to that of the planktonic embryos of sea urchins and different from that of embryos developing from terrestrial eggs, freshwater eggs, and planktonic and demersal marine eggs.Although broods varied widely in egg number and mean egg dry weight, C. curata gives eggs which contain a constant proportion of organic components.Levels of ash, water, and protein in the adults exceeded those in the pentacula, and lipid comprises a much smaller proportion of the adult body than it did of the pentacula.     

Function of shell structures of pig louse and how egg maintains a low equilibrium temperature in direct sunlight

The functional significance of the structures of the eggshell of the pig louse, Haematopinus suis, and those of some other Anoplura and some Mallophaga, is described.The shell of the pig louse has an outer air-filled meshwork. The air in this meshwork is isolated from that in the respiratory system. This layer does not function as an insulator. Calculations show that the temperature drop across the layer is only about $\text{1}\text{75}\text{°}\text{C}$.The air-filled outer meshwork functions as a reflector. The eggs are often exposed to direct sunlight because of the wide spacing of the pig's bristles. The outer reflecting layer has an enormous selective advantage for the egg in two quite different ways: (1) it slows down the rate at which equilibrium temperature is reached, and (2), much more important, it enables the egg to lower equilibrium temperature.Assuming that reflectance = 0.9 and sunlight = 10³ Wm^?2, the heat input is $\text{3.1 ×}\text{r}{}^2\text{100}\text{J/sec}$, and the time required for a rise of 20°C, assuming no heat loss, is 4.3 min. Assuming a more realistic energy flux (sunlight = 500 Wm^?2) for the latitudes in which most pigs are bred, and even assuming a reflectance of only 20%, the time required for a rise of 20°C in direct sunlight, with no heat loss, is increased by 1 min 4 sec.Lowering the equilibrium temperature is of much greater selective advantage than merely slowing down the rate at which it is reached. Reflectance + emissivity = 1. Nevertheless, the egg can achieve a reflectance of, say, 0.8 at the same time as it achieves an emissivity of 0.8 from the energy absorbed by the chorion and especially the aqueous embryo or pharate first instar larva, either of which will function as a black body and absorb all of the infra-red not absorbed by the chorion. Calculations show that if a heat input of 200 Wm^?2 is assumed, at an ambient temperature of 27°C, the equilibrium temperature can be maintained at 47°C.The chorionic hydropyle of the pig louse, like that of apparently all other Phthiraptera, is most unusual in that the hygroscopic spumaline secreted by the accessory glands flows through the hydropyle canals of the chorion into the interior of the egg while the egg is still in the common oviduct. Molecules as large as those of fast green (mol. wt 809) pass through the spumaline into the interior of the egg and then pass through the serosal hydropyle. The amount of spumaline secreted greatly exceeds that necessary to glue the egg to the bristle. Once absorption is seen as another function of the spumaline, its amount no longer becomes difficult to understand: under many natural conditions it will enable the egg quickly to replace losses of water that result from exposure to high temperatures in direct sunlight.The egg of the pig louse, like those of other species of Haematopinus, has a very well-developed plastron or permanent physical gill. A number of the aeropyles are covered by spumaline, but even assuming that 30% of them are so covered, the water-air interface for small eggs is 3.5 × 10⁶ μm²/mg and for large eggs 2.6 × 10⁶ μm²/mg. Thus the relation of water-air interface to weight of tissue is the most favourable so far reported in insects.     

L'oeuf de Myrmeleotettix maculatus Thunb. (Orthoptera : Acrididae): Absorption d'eau et structure fine des enveloppes

Water absorption and fine structure of the egg shell have been studied in the eggs of Myrmeleotettix maculatus (Orthoptera : Acrididae). At the anterior pole of these eggs, there is a respiratory plastron and at the posterior end, behind the ring of micropyles, many minute holes can be seen. Eggs of M. maculatus incubated on wet filter paper at 30°C absorb water before diapause between the 5th and the 25th day after oviposition. The period of water absorption coincides with the serosal cuticle secretion. The increase in volume of the egg does not seem to involve any fragmentation in the chorion. The chorion is composed of 3 layers (upper, middle and lower). The middle layer, the thickest one, contains twisted elements with a paracrystalline structure. During the first days of incubation, the vitelline membrane becomes changed. In the newly laid egg, it is granular and homogeneous. Five days later, 4 layers with different structures can be distinguished. After 30 days, the serosal cuticle presents a thin epicuticule and a thick lamellated endocuticule with many pore canals.     

Comparison of the physical properties of real and artificial salmonid eggs and of their performance when drifting in an experimental stream channel

The paper compares three batches of brown trout (Salmo trutta Linnaeus) eggs, two batches of Atlantic salmon (Salmo salar Linnaeus) eggs and the artificial trout eggs described by Ottaway (1981), in terms of their measurable physical characteristics. Attention was given to the effects of temperature and of the stage of development of the real eggs. Comparisons were also made between brown trout eggs and artificial eggs of the rate of fall in a water column, of the value and speed of attainment of terminal velocity when falling through water and of the pattern of settlement within an experimental stream channel.The main measurable characteristics of brown trout eggs (after water hardening) and of artificial eggs (values in parentheses) were: diameter 0.52–0.54 mm (0.54), fresh weight 0.078–0.086 g (0.09), density 1.071–1.075 g ml^–1 (1.066), and volume 0.073–0.081 ml (0.085). There was no evidence of any major change in these values as egg development proceeded. Salmon eggs had a similar density to trout and artificial eggs but had 30 to 70% greater weight and volume. The water-hardening of eggs caused their dry matter content to fall from 37–39% to 31–34% and there were corresponding increases in volume and fresh weight.The rate of fall of trout eggs and artificial eggs through water when timed from rest at the water surface to a depth of 125 cm was similar for both types of egg at 10 °C. This rate rose with increasing temperature at the same low rate of c. 0.02 cm s^–1 °C^–1 for both types of egg. Both types of egg had a terminal velocity of c. 8.8 cm s^–1 and both achieved terminal velocity in less than 1.5 s after release.The patterns of settlement of the two types of eggs in an experimental channel were similar.     

Analysis of some morphological and biochemical characteristics of the egg of the predaceous bugMacrolophus caliginosus (Het.: Miridae) during embryogenesis

The predaceous bug,Macrolophus caliginosus Wagner (Heteroptera: Miridae) is a biological control agent used in greenhouses in the French Mediterranean region. The egg of this mirid is embedded in plant tissues and all embryonic development occurs in this environment. In order to define an artificial substrate for egg-laying, exchanges of chemical substances between the egg and the host plant were investigated by morphological and biochemical analyses. Egg wet weight doubled throughout embryonic development, while egg dry weight did not vary. The amino acid content of the egg remained stable during embryogenesis. Only aspartic acid, serine and alanine decreased during the first five days of development. It can be concluded that exchanges mainly involved uptake of water by the egg.     

Entomopathogenic Nematodes and Bacteria Applications for Control of the Pecan Root-Knot Nematode, Meloidogyne partityla, in the Greenhouse

Meloidogyne partityla is a parasite of pecan and walnut. Our objective was to determine interactions between the entomopathogenic nematode-bacterium complex and M. partityla. Specifically, we investigated suppressive effects of Steinernema feltiae (strain SN) and S. riobrave (strain 7–12) applied as infective juveniles and in infected host insects, as well as application of S. feltiae''s bacterial symbiont Xenorhabdus bovienii on M. partityla. In two separate greenhouse trials, the treatments were applied to pecan seedlings that were simultaneously infested with M. partityla eggs; controls received only water and M. partityla eggs. Additionally, all treatment applications were re-applied (without M. partityla eggs) two months later. Four months after initial treatment, plants were assessed for number of galls per root system, number of egg masses per root system, number of eggs per root system, number of eggs per egg mass, number of eggs per gram dry root weight, dry shoot weight, and final population density of M. partityla second-stage juveniles (J2). In the first trial, the number of egg masses per plant was lower in the S. riobrave-infected host treatment than in the control (by approximately 18%). In the second trial, dry root weight was higher in the S. feltiae-infected host treatment than in the control (approximately 80% increase). No other treatment effects were detected. The marginal and inconsistent effects observed in our experiments indicate that the treatments we applied are not sufficient for controlling M. partityla.     

Artificial incubation of California condor Gymnogyps californianus eggs removed from the wild

The currrent California condor (Gymnogyps californianus) recovery plan entails increasing the reproductive rate via replacement-clutch manipulation of eggs. During the period from 1983 to 1985, 15 eggs were removed from wild nesting pairs for artificial incubation. The eggs were incubated at a dry bulb temperature of 36.4°C in modified forced-air Lyon Electric incubators. The incubation humidity was adjusted for individual eggs based on weight loss data (water = weight), 25.6–30.0°C wet bulb (41.0–63.0% Relative Humidity (RH)). The chicks were hatched initially under forced-air conditions of 36.1°C dry bulb, 31.1–01.7°C wet bulb (70.0–73.0% RH). In 1984, hatching parameters were changed to still-air conditions, 36.1°C dry bulb (top of the egg), 35.0°C dry bulb (bottom of the egg), 31.1–31.7°C wet bulb (70.0-73.0% RH). Tactile and auditory stimulation was utilized during the pip-to-hatch interval. From among 15 eggs collected, 13 hatched, and 12 condor chicks were raised successfully (hatchability: 86.7%; survivability: 92.3%).     

Biometrical relationships in developing eggs and neonates of Octopus vulgaris in relation to parental diet

Captive Octopus vulgaris adults were fed three mono-diets based on pilchard, crab and squid and allowed to grow until reproduction under controlled temperature. Spawns from each dietary treatment were isolated, and the embryonic development, egg length, width and wet weight, in addition to neonate dry weight, dorsal mantle length and ventral mantle length were monitored. Pilchard-diet spawns developed faster in terms of thermal time. Initial egg wet weight was higher for squid and crab diets. Irrespective of the parental diet, eggs passed through a swelling process so that egg width and wet weight increased in a nonlinear way, whereas egg length was left nearly unaffected. Egg length and initial wet weight showed a high correlation with neonate dry weight. Egg length, even at advanced incubation, can be used as a good proxy for neonate dry weight, this fact having potential implications for the ecological and aquaculture research on O. vulgaris.     

STUDIES ON CELL METABOLISM AND CELL DIVISION : VI. OBSERVATIONS ON THE GLYCOGEN CONTENT,CARBOHYDRATE CONSUMPTION,LACTIC ACID PRODUCTION,AND AMMONIA PRODUCTION OF EGGS OF ARBACIA PUNCTULATA

1. Under the present conditions of experiment, Arbacia eggs were found to contain an average of 110 mg. of acid-hydrolyzable carbohydrate (calculated as glucose) per gm. of egg protein. This carbohydrate was almost all in the egg proper, little or none being found in the jelly. To permit conversion of the data to other bases of reference the relation of nitrogen content to wet and dry weight and to egg number were determined. The eggs were found to contain 23.9 per cent solids, 0.10 mg. nitrogen per mg. dry weight, and 5.93 mg. nitrogen per 10⁶ cells. From these results, about 7 per cent of the egg dry weight is acid-hydrolyzable carbohydrate and about 65 per cent is protein. 2. Approximately one-half of the total acid-hydrolyzable carbohydrate was isolated in the form of an alkali-stable, alcohol-precipitable carbohydrate. This substance gave a typical glycogen color test with iodine, yielded glucose on acid hydrolysis, and had, within the limits of experimental error, the same optical rotation as glycogen from other animal sources. Since known amounts of glycogen were completely recovered when carried through the isolation process, the nature of one-half of the acid-hydrolyzable carbohydrate of Arbacia eggs remains undetermined. 3. In order to gain some estimate of the extent to which Arbacia eggs utilize their total carbohydrate for development, determinations of the oxygen consumption, respiratory quotient, carbohydrate consumption, lactic acid production, and ammonia production were made. While all samples of eggs were found to utilize carbohydrate from the 15th to the 24th hours of development at 20°C., certain samples of eggs consumed little or no carbohydrate from the 1st to the 6th hours, the period during which cell division proceeds most rapidly. In a number of instances where carbohydrate breakdown was lacking, a substantial proportion of the oxygen consumption could be accounted for on the basis of processes involving oxidation of protein or protein breakdown products.     

Ultrastructure de l'œuf de Triatoma infestans Klug

Summary The thick rigid chorion of the egg of Triatoma secreted by the follicle cells shows two porous layers: an aerial layer in the exochorion, an alveolar one in the endochorion. The anterior part of the eggshell is closed up by an operculum which is heaved up by the hatching larva. The operculum has no alveolar layer. The air enters through the numerous holes of the shell surface into the aerial layer and through the micropyles into the alveolar layer. The egg has no respiratory plastron.The follicle cells produce also a vitelline envelope whose structure shows a rapid condensation at fertilization time. During its development the embryo secretes two layers: serosal and embryonic cuticle.At high humidities, at low temperatures the egg is able to increase its weight during the early stages of embryogenesis, and this increase stops when the serosal cuticle is secreted. In a dry atmosphere the egg loses water but can develop if the temperature is higher than 20°C.The little permeability of the egg is related to the structure of its envelopes. The chorion and the vitelline envelope prevent the water from getting out of the egg. The serosal cuticle seems to be opposed to the penetration of the water into the egg. The role of the embryonic cuticle is probably limited in the transit of water.

Nous remercions Messieurs les Professeurs Maillet et Folliot qui ont mis le microscope R.C.A. à notre disposition, Madame Allo et Mademoiselle Le Gac, technicienne au microscope à balayage J.S.M. S1, pour leur collaboration technique.     

Carbohydrate metabolism and restricted oxygen supply in the eggs of the silkworm, Bombyx mori

Increased oxygen supply to diapause eggs of the silkworm (O₂-incubation) effectively prevented diapause initiation and induced the same pattern of glycogen, polyol and lactate levels as was observed in normal non-diapause eggs. Sensitivity to oxygen decreased as embryonic development proceeded. After the termination of this sensitive period, accumulation of polyols and lactate followed.Experiments were carried out to test whether changes in the oxygen permeability of the egg membranes are involved in restricting the supply of this gas to eggs at the onset of diapause. Oxygen permeability of the chorion was measured with apparatus especially designed for this purpose. Although the chorion of the diapause egg was less permeable than that of the non-diapause egg, the oxygen permeability of the chorion does not change appreciably during the early developmental stages of the diapause eggs. The changes in rate of water loss through the egg membranes were measured during the early developmental stages of the embryos. The level of water loss decreased gradually as the formation of serosal cuticle proceeded. Moreover, it was observed that the water loss up to the time of formation of serosal cuticle was closely related to the oxygen permeability of the chorion.From these results, we suggest that the formation of the serosal cuticle may be an additional cause of the restricted oxygen supply at the onset of the diapause.     

Physiological changes during colony establishment in the termite Hodotermes mossambicus (Hagen): Water balance and energy content

The changes in dry mass, wet mass and energy content were determined during the establishment of the incipient colony. The winged reproductives of H. mossambicus are weak fliers. It is postulated that the alates compensate for the large quantity of reserve material by reducing their body water. The wet mass of both males and females increased significantly (twice original mass) from emergence until the first larvae appeared which tends support to this view. Furthermore, although the termites maintained in groups imbibed water, it is also clear that the reproductives, as long as they remain social, maintain the same low body-water content. This desiccated physiological condition appears to be and adaptation which assists flight and dispersal. In contrast with sterile eggs, fertile eggs absorbed water during development. The paired females (normal and homosexual) maintained the same energy/unit wet mass from egg production until the first workers appeared. Thus the possibility exists that the hydrophobic reserves (e.g. triglycerides) were converted to the hydrophilic reserves (e.g. carbohydrates). The same applied to the eggs and offspring.The changes in dry mass, wet mass, body water and energy content of females per unit of eggs (specific mass) produced were 0.005, 8.696, 21,276 and 12.820 respectively. The small specific dry mass value can probably be ascribed to the use of small amounts of glycerides and the increase in dry mass by the conversion from relatively light and “compact” glycerides to relatively heavy and “bulky” glycogen.Females utilised far more dry mass and energy in comparison to males to survive until the first workers appeared. Wet mass, body water and energy used in rearing of the offspring shows clearly that males made a larger contribution to the water and energy requirements of the offspring.     

Changes in lipid and fatty acid composition of eggs during development of the beet armyworm, Spodoptera exigua

The dry weight of Spodoptera exigua eggs decreased by 15·9 μg/egg or 64% of initial weight during embryogenesis and development of pharate first instar larvae. Lipid depletion accounts for 36% of this total dry weight loss and this occurs at an essentially constant rate throughout development. This marks S. exigua as an exception since most insects utilize lipids more rapidly during later developmental stages. Lipid depletion is due primarily to triglyceride catabolism, although phospholipids also decrease significantly.Fatty acid composition remains stable during development. In triglycerides, 18:1 is most common followed by 16:0 and 18:2; in the phospholipids, the order of abundance is 18:1, 18:2, and 16:0. Egg fatty acids differ from dietary fatty acids: 16:1 comprises 7% of triglyceride fatty acids although it is not present in the larval media; 18:1 predominates in the egg whereas 18:2 is most abundant in the diet.     

The role of pressure in controlling the entry of water into the developing eggs of the Australian plague locust Chortoicetes terminifera (Walker)

The normal internal hydrostatic pressure and the additional pressure necessary to rupture the egg shell was measured in the eggs of Chortoicetes terminifera, Newly laid white eggs burst at c 0.15 kg cm^-2, but after external tanning the chorion withstands c 0.5 kg cm^-2 when removed from its tanned foam ‘corset’ and 1.0 kg cm^-2 if left embedded in the egg pod material. Older eggs with formed cuticles often withstand 2.0 kg cm^-2 but yield at rather lower pressures if they develop ‘pin-holes’. As the OP of the egg contents always exceeds 7.7 kg cm^-2 the rigidity of the wall is clearly insufficient to permit the generation of high hydrostatic pressures capable of preventing water entry during the non-absorbing phases of development. Real hydrostatic pressures are lower than 0.06 kg cm^-2 in the young intact egg and reach only c 0.5 and 0.3 kg cm^-2, respectively, during the absorptive and post-absorptive phases of development. Several events contribute to the sigmoid form of the water uptake curve. Water is at first excluded by a permeability barrier associated with the chorion. Absorption is delayed until the yolk is completely enclosed by the serosal cell layer. After undergoing cleavage, the yolk is then rapidly mobilized to furnish precursors for cuticle synthesis; in consequence, the internal OP rises from δ 0.76d?K to 0.93d?K despite the massive inflow of water which is governed by the osmotic gradient. At blastokinesis the serosa becomes detached from the cuticle; cuticle deposition and yolk mobilization are halted, the OP falling rapidly to cδT 0.53d?K. The bulk entry of water then ceases. Any excessive hydrostatic pressures which develop later are relieved by the formation of self-sealing ‘pin-holes’.     

A study on the energy source in the developing embryo of the mangrove horseshoe crab,Carcinoscorpius rotundicauda (Latreille)

The mangrove horseshoe crab, Carcinoscorpius rotundicauda, is an Asian species found in Malaysia. This ancient arthropod has a long incubational period during which it depends on various energy sources for both embryogenesis and organogenesis. This study describes the trend of energy utilization from the endogenous reserves by the developing embryos from 0 to 40 days of incubation (until the hatching of the larvae). The dry weight, insoluble protein, carbohydrate, glycogen and lipid showed a declining trend from 0 to 40 days of incubation, whereas the wet weight, water content, ash content and soluble protein showed an increasing trend. Selected micro-elements such as Cu²⁺, Fe²⁺ and Zn²⁺ also demonstrated an interesting trend in the developing eggs when the egg mass was subjected to inductively coupled plasma mass spectrometry analysis, where these elements showed a high correlation with the moulting stages and egg development. Maximum variations within the micro-elements were observed during the 1st (10 days after fertilization) and 2nd (35–36 days after fertilization) moulting stages within the developing eggs. This study clearly indicated that the moulting cycles of C. rotundicauda during embryonic development influence energy utilization in the form of carbohydrates, lipids, proteins, glycogen and other micro-elements.     

Long-term droughts change the hatching patterns of zooplankton resting eggs from permanent and temporary lakes

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STUDIES ON CELL METABOLISM AND CELL DIVISION : VIII. THE DIPHOSPHOPYRIDINE NUCLEOTIDE (COZYMASE) CONTENT OF EGGS OF ARBACIA PUNCTULATA

1. The diphosphopyridine nucleotide content of Arbacia eggs has been measured manometrically and found to be approximately 250–500 micrograms per gm. wet weight of eggs, the value varying with individual egg samples and with the state of development of the eggs. Of the total diphosphopyridine nucleotide present, approximately 25–40 per cent is in an alkali-stable, presumably the dihydro, form. 2. Tests for triosephosphate and glutamic acid dehydrogenases carried out on Arbacia egg cytolysates were negative.