Effects of experimental manipulation of inflorescence size on pollination and pre-dispersal seed predation in the hummingbird-pollinated plant Ipomopsis aggregata

Large floral displays should theoretically provide advantages to plants through increased pollinator visitation and resulting fruit and seed set. However empirical tests of the response of pollinators to floral display size have been limited by a lack of direct experimentation, and the results of such studies have been equivocal. In addition, other selective agents such as pre-dispersal seed predators might modulate effects of floral display on pollination. By artificially altering flower number, we examined the direct effects of floral display in the monocarpic herb, Ipomopsis aggregata (Polemoniaceae), on visitation rates by broad-tailed and rufous hummingbird pollinators, as well destruction of fruits by a pre-dispersal seed predator (Hylemya: Anthomyiidae). In addition, we quantified the ultimate effects of flower number on female reproductive success. Plants with larger floral displays were most likely to be visited first in any given foraging bout (P < 0.01). As expected, plants with more flowers received more total flower visits. However, we found no gain in the proportion of flowers visited for many- versus few-flowered plants, or the total number of approaches/hour. In fact, a significantly greater percentage of flowers were visited on few-flowered plants. Plants did not compensate for our reduction in flowers by increasing investment in the number or proportion of flowers that set fruit, the number of seeds/fruit, or seed weight. Pre-dispersal seed predation was greater for many- than for few-flowered plants (P < 0.001), but this did not offset the potential fitness gains of producing large displays. Our data support the hypothesis that large floral displays function primarily in long-distance attraction of pollinators, and enhance maternal success. Received: 21 March 1996 / Accepted: 24 October 1996     

The birch tube-maker Acrobasis betulella in a fragmented habitat: the importance of patch isolation and edges

We examined the response of the birch tube-maker Acrobasis betulella Hulst (Lepidoptera: Pyralidae) to habitat patch isolation and edges. Density of A.␣betulella larva was higher on distant islands than on islands close to the shore. Following experimental removal of all larvae from the islands, adults were able to recolonize even the most distant islands, and larval density was again positively correlated with the degree of isolation. Larval density was not correlated with island size or the amount of birch present on the islands. Larvae on more distant islands did not have lower mortality than those in less isolated sites. Larvae were found more often on edges than in the interior of birch stands and developed faster on edges. This positive edge effect, coupled with the ability of the adults to disperse to even our most distant islands, suggests that A. betulella would be favored in an environment fragmented at the same scale as our island system. Received: 15 April 1996 / Accepted: 30 September 1996     

Effects of harvester ants on plant species distribution and abundance in a serpentine grassland

Seed harvesting ants can have important effects on the composition and structure of plant communities. We investigated two effects of Messor andrei, the black seed-harvesting ant, on a serpentine grassland plant community in northern California. First, to determine if selective seed predation by ants affects plant community composition, we excluded harvester ants from 1-mediameter circular plots of grassland. Abundances of all species on these plots and on control plots were measured before and after exclosure. Second, to determine if M. andrei nest mounds affect plant community composition, we compared plant species abundances on and off nest mounds. M. andrei deposit large amounts of organic matter on their nest mounds over a foraging season, so mounds may alter the edaphic environment. The exclusion of seed-harvesting activity did not cause changes in the plant community. Nest mounds had a strong effect on plant communities: there were many more grasses and fewer forbs on ant mounds, although at least one forb, Lepidium nitidum, produced twice as many seeds when it grew on nest mounds. We found that nest mounds formed islands of higher-temperature soil in the serpentine grassland. Received: 31 March 1997 / Accepted: 6 May 1997     

Effects of 3 days of carbohydrate supplementation on muscle glycogen content and utilisation during a 1-h cycling performance

This study compared the effects of supplementing the normal diets of six trained cyclists [maximal oxygen uptake O_2max) 4.5 (0.36)l · min⁻¹; values are mean (SD)] with additional carbohydrate (CHO) on muscle glycogen utilisation during a 1-h cycle time-trial (TT). Using a randomised crossover design, subjects consumed either their normal diet (NORM) for 3 days, which consisted of 426 (137) g · day⁻¹ CHO [5.9 (1.4) g · kg⁻¹ body mass (BM)], or additional CHO (SUPP) to increase their intake to 661 (76) g · day⁻¹ [9.3 (0.7) g · kg⁻¹ BM]. The SUPP diet elevated muscle glycogen content from 459 (83) to 565 (62) mmol · kg⁻¹ dry weight (d.w.) (P < 0.05). However, despite the increased pre-exercise muscle glycogen stores, there was no difference in the distance cycled during the TT [40.41 (1.44) vs 40.18 (1.76) km for NORM and SUPP, respectively]. With NORM, muscle glycogen declined from 459 (83) to 175 (64) mmol · kg⁻¹ d.w., whereas with SUPP the corresponding values were 565 (62) and 292 (113) mmol · kg⁻¹ d.w. Accordingly, both muscle glycogen utilisation [277 (64) vs 273 (114) mmol · kg⁻¹ d.w.] and total CHO oxidation [169 (20) vs 165 (30) g · h⁻¹ for NORM and SUPP, respectively] were similar. Neither were there any differences in plasma glucose or lactate concentrations during the two experimental trials. Plasma glucose concentration averaged 5.5 (0.5) and 5.6 (0.6) mmol · l⁻¹, while plasma lactate concentration averaged 4.4 (1.9) and 4.4 (2.3) mmol · l⁻¹ for NORM and SUPP, respectively. The results of this study show that when well-trained subjects increase the CHO content of their diet for 3 days from 6 to 9 g · kg⁻¹ BM there is only a modest increase in muscle glycogen content. Since supplementary CHO did not improve TT performance, we conclude that additional CHO provides no benefit to performance for athletes who compete in intense, continuous events lasting 1 h. Furthermore, the substantial muscle CHO reserves observed at the termination of exercise indicate that whole-muscle glycogen depletion does not determine fatigue at this exercise intensity and duration. Accepted: 25 November 1996     

Effect of seed predation on seed bank size and seedling recruitment of bush lupine (Lupinus arboreus)

Whether seed consumers affect plant establishment is an important unresolved question in plant population biology. Seed consumption is ubiquitous; at issue is whether seedling recruitment is limited by safe-sites or seeds. If most seeds inhabit sites unsuitable for germination, post-dispersal seed consumption primarily removes seeds that would otherwise never contribute to the population and granivory has minimal impacts on plant abundance. Alternatively, if most seeds ultimately germinate before they lose viability, there is greater potential for seed consumption to affect plant recruitment. Of the many studies on seed consumption, few ask how seed loss affects seedling recruitment for species with long-lived seed banks. We examined post-dispersal seed predation and seedling emergence in bush lupine (Lupinus arboreus), a woody leguminous shrub of coastal grasslands and dunes in California. We followed the fate of seeds in paired experimental seed plots that were either protected or exposed to rodent granivores in grassland and dune habitats. Significantly more seeds were removed by rodents in dunes than grasslands. In dunes, where rodent granivory was greatest (65% and 86% of seeds removed from plots by rodents in two successive years), there is a sparse seed bank (6.6 seeds m⁻²), and granivory significantly reduced seedling emergence (in the same two years, 18% and 19.4% fewer seedlings emerged from exposed versus protected plots), suggesting seed rather than safe-site limited seedling recruitment. In contrast, rodents removed an average of 6% and 56% of seeds from grassland plots during the same two years, and the grassland seed bank is 43-fold that of the dunes (288 seeds m⁻²). Even high seed consumption in the second year of the study only marginally influenced recruitment because seeds that escaped predation remained dormant. Burial of seeds in both habitats significantly reduced the percentage of seeds removed by rodents. Results suggest that granivores exert strong but habitat-dependent effects on lupine seed survival and seedling emergence. Received: 24 October 1996 / Accepted: 4 February 1997     

Parental quality and egg size affect chick survival in a precocial bird, the lapwing Vanellus vanellus

Effects of egg size and parental quality on lapwing Vanellus vanellus chick survival were studied in southwestern Sweden over 6 years. Chicks from large eggs were heavier at hatching and survived significantly better than those from small eggs. To control for the confounding effect of parental quality on egg size and chick survival, we performed a cross-fostering experiment during 2 years, exchanging clutches between nests with large and small eggs. In control clutches, chicks from large eggs survived better than those from small eggs, but we found no significant difference in chick survival between exchanged clutches. Thus, egg size did not affect chick survival independently of parental quality. Fledging success increased with parental age and/or experience, and with female body mass. Hence, both egg size and parental quality affect chick survival in the lapwing. Received: 22 February 1996 / Accepted: 30 September 1996     

The impact of vertebrate and invertebrate predators on a stream benthic community

I assessed the impact of both vertebrate and invertebrate predators on a lotic benthic community in a 1-month-long experiment, using enclosures containing cobble/gravel bottoms, with large-mesh netting that allowed invertebrates to drift freely. Brown trout (Salmo trutta) and leeches (Erpobdella octoculata) were used as predators and four treatments were tested: a predator-free control, leeches only, trout only, and leeches and trout together. A density of 26.7 leeches/m² (20 leeches/enclosure) and 1.3 trout/m² (one trout per enclosure) was stocked into the enclosures. The total biomass of invertebrate prey was significantly lower in the trout and trout plus leech treatments than in the leech and control treatments, which were due to strong negative effects of trout on Gammarus. On the individual prey taxon level, both trout and leeches affected the abundance of Asellus , Baetis and Ephemerella, whereas the abundance of Gammarus was only affected by trout, and the abundance of Orthocladiinae and Limnephilidae was only affected by leeches. In the treatment with trout and leeches together, the abundance of Ephemerella and Baetis was higher than when trout or leeches were alone, which was probably due to predator interactions. Leeches and trout had no effects on prey immigration but did affect per capita emigration rates. Both trout and leeches indirectly increased periphyton biomass in enclosures, probably due to their strong effects on grazers. Both trout and leeches were size-selective predators, with trout selecting large prey, and leeches selecting small prey. Size-selective predation by trout and leeches affected the size structure of five commonly consumed prey taxa. Trout produced prey populations of small sizes owing to consumption of large prey as well as increased emigration out of enclosures by these large prey. Leech predation produced prey assemblages of larger size owing to consumption and increased emigration of small prey. These results suggest that in lotic habits, predatory invertebrates can be as strong interactors as vertebrate predators. Received: 23 June 1997 / Accepted: 4 May 1998     

Effects of hydration state on hormonal and renal responses during moderate exercise in the heat

The effects of hydromineral hormones and catecholamines on renal concentrating ability at different hydration states were examined in five male volunteers while they performed three trials. Each of these trials comprised a 60-min exercise bout on a treadmill (at 50% of maximal oxygen uptake) in a warm environment (dry bulb temperature, 35°C; relative humidity, 20–30%). In one session, subjects were euhydrated before exercise (C). In the two other sessions, after thermal dehydration (loss of 3% body mass) which markedly reduced plasma volume (PV) and increased plasma osmolality (osm_pl), the subjects exercised either not rehydrated (Dh) or rehydrated (Rh) by drinking 600 ml of mineral water before and 40 min after the onset of exercise. During exercise in the Dh compared to C state, plasma renin, aldosterone, arginine vasopressin (AVP), noradrenaline and adrenaline concentrations were increased (P < 0.05). A reduction in creatinine clearance and urine flow was also observed (P < 0.05) together with a decrease in urine osmolality, osmolar clearance and sodium excretion, while free water clearance increased (P < 0.05). However, compared to Dh, Rh partially restored PV and osm_pl and induced a marked reduction in the time courses of both the plasma AVP and catecholamine responses (P < 0.05). Values for renal water and electrolyte excretion were intermediate between those of Dh and C. Plasma atrial natriuretic peptide presented similar changes whatever the hydration state. These results demonstrate that during moderate exercise in the heat, renal concentrating ability is paradoxically reduced by prior dehydration in spite of high plasma AVP levels, and might be the result of marked activation of the sympatho-adrenal system. Rehydration, by reducing this activation, could partially restore the renal concentrating ability despite the lowered plasma AVP. Accepted: 23 April 1997     

Effects of ultraviolet radiation on early larval stages of the Alpine newt, Triturus alpestris, under natural and laboratory conditions

Although Alpine newts (Triturus alpestris) are found at altitudes up to 2500 m, their larvae proved to be extremely sensitive to UV radiation when exposed in clear tapwater to natural sunlight or to comparable artificial UV-B radiation in the laboratory. The experiments revealed severe skin damages (lysis of epithelial cells) and mortality after a few days of exposure. In their natural habitats above the timberline, however, the larvae are protected by the high concentration of dissolved organic carbon in the water leading to an almost complete absorption of UV radiation within the first few centimeters of the water layer. Furthermore, when exposed to UV radiation larvae show erratic swimming activities that may carry them into protected areas. Only in very shallow natural habitats did we detect sublethal UV-like histological effects. Shallow high mountain ponds with clear water normally lack newt populations, probably as a consequence of both low temperature and the effects of UV radiation. Received: 4 October 1996 / Accepted: 7 February 1997     

The effect of light environment, leaf area, and stored carbohydrates on inflorescence production by a rain forest understory palm

Variation in flowering by long-lived plants may be correlated with current resource availability. If, however, there are trade-offs between current and future reproduction, or between reproduction and storage or growth, then understanding variation requires a whole-plant, longer-term perspective. Inflorescence production by Calyptrogyne ghiesbreghtiana Linden ex. H. Wendl., an understory palm, was studied over 3 years. Annual inflorescence production varied greatly and was correlated with variation in plant size and light environment. There was no trade-off between past inflorescence production and the frequency of future inflorescence production. On the contrary, individuals that produced more inflorescences than predicted from their size and light environment tended to continue to do so in subsequent years also. I manipulated the resource environment of a subset of plants by removal of leaves and/or reproductive spikes. Leaf removal suppressed inflorescence production for the following 2 years, but spike removal had no effect. One year after leaf removal stored reserves were, on average, back to pre-treatment levels. There was, however, a negative effect of recent inflorescence production on storage. Plants with higher levels of storage had higher inflorescence production in the next 75 days. In C. ghiesbreghtiana the resource cost of reproduction is apparent in short-term variation in stored reserves. In contrast, annual inflorescence production does not follow a trade-off pattern between successive years, but consistently reflects both plant size and the light environment. Received: 20 October 1996 / Accepted: 25 January 1997     

The influence of pollen addition and ramet isolation on current sexual reproduction in a clonal herb

Canada mayflower (Maianthemum canadense Desf.), a rhizomatous, perennial herb, was the subject of a 2-year field experiment that examined two factors potentially affecting fruit and seed production: pollen addition and ramet isolation. Ramets were either open pollinated or overpollinated by hand to supplement natural levels. Rhizomes of the ramets were either severed, to prevent resource supplementation from the genet, or left intact. Ramets that were overpollinated matured more fruits and more seeds than ramets that were open pollinated. Thus, mayflower appears to have been pollen limited in both years. Ramets that were open pollinated and whose rhizomes were severed matured as many fruits, seeds, and seeds/fruit as ramets that were open pollinated and whose rhizomes were left intact. Ramets that were overpollinated and whose rhizomes were severed matured fewer fruits, seeds, and seeds/fruit than overpollinated ramets whose rhizomes were left intact. It appears that at natural levels of pollination, mayflower ramets are physiologically independent but as the level of pollen increases, mayflower ramets receive support from other parts of the genet. Received: 13 May 1996 / Accepted: 1 November 1996     

The coadaptation of female morphology and offspring size: a comparative analysis in crickets

Few studies of invertebrates have considered combinations of morphological and life history traits in the context of the evolution of reproductive strategies. Cricket species that exploit habitats harsh with respect to egg survival have evolved a long ovipositor, presumably because laying deep in the soil reduces egg mortality. Yet hatchling mortality increases with laying depth, and the ability of hatchlings to climb through the soil increases with egg size. Thus a conflict may exist between survival of the egg and that of the hatchling, inducing a positive covariation between egg size and ovipositor length across species evolving under contrasting selective habitats. We used the phylogenetic autocorrelation method and a path analysis to assess whether egg size coevolved with ovipositor length across 40 species of crickets, and whether egg size was affected by body size or ecological factors that influence egg mortality. Body size and ovipositor length were affected by taxonomic association, whereas common ancestry had no significant effect on egg size, diapausing strategy, and oviposition preference for soil types. The path model indicated that 29.11% of the variance in egg size was explained by independent evolution. As expected, ovipositor length was positively correlated with egg size, and species diapausing in the egg stage produced larger eggs than crickets diapausing in the nymphal stage or with no diapause. Ovipositor length and diapausing strategy were the first and second most important traits, respectively, in term of the proportion of variance in egg size explained by specific values. These results support the hypothesis that the ability of hatchlings to climb through the soil, and variation in diapause strategies, are general selective factors affecting the evolution of egg size in crickets. Phylogeny explained 51.01% of the variance in egg size. Egg size in a current cricket species, however, was not directly determined by egg size in its ancestor. Instead, it was strongly related to the phylogenetic values of body size and ovipositor length. Such indirect phylogenetic effects of body size and ovipositor length may have arisen because clades originating from ancestors with different ovipositor lengths experienced different selective pressures on egg size. Recelived: 13 October 1995 / Accepted: 30 September 1996     

Incubation capacity and clutch size determination in two calidrine sandpipers: a test of the four-egg threshold

Several groups of vertebrate taxa, including shorebirds, are unusual in that they produce a fixed number of offspring. The aim of this study was to examine whether the incubation capacity of western sandpipers (Calidris mauri) and semipalmated sandpipers (C. pusilla) limits their maximum clutch size to four eggs. Experimental enlargement of clutch size had no effect on rates of nest abandonment, nest attendance or loss of body mass by incubating sandpipers. The duration of incubation was significantly longer for enlarged five-egg nests, and there were trends towards increased partial clutch loss and asynchrony at hatch, but overall hatching success was unaffected by experimental egg number. I conclude that small, calidrine sandpipers with biparental care are able to compensate for an additional egg in an enlarged nestbowl, despite the constraints of conically shaped eggs and two brood patches. Possibly, shorebirds do not lay more than a fixed clutch size of four eggs because selection on factors acting during egg production or brood-rearing is more important in regulating offspring number. Received: 20 June 1996 / Accepted: 30 September 1996     

Effects of seed size and emergence time on tree seedling establishment: importance of developmental constraints

Arafura filesnakes (Acrochordus arafurae) are large (up to 2.5 m, 5 kg) aquatic nonvenomous snakes that feed entirely on fishes. A 10-year field study in the Australian wet-dry tropics revealed strong correlations between rainfall patterns, fish abundance, and snake population dynamics. All of these characteristics showed considerable annual variation. High rainfall late in the wet season (February–March) caused prolonged inundation of the floodplain. Following such years, dry-season sampling revealed that fishes were abundant, filesnakes were in good body condition, and a high proportion of adult female filesnakes were reproductive. Annual variation in recruitment to the population (as judged by the relative abundance of yearling snakes) was also correlated with fish abundance and thus, with rainfall patterns in the late-wet season. Our results fit well with those from other studies on a diverse array of aquatic and terrestrial species within the wet-dry tropics. Annual variation in rainfall patterns, via its effects on prey abundance, may drive the population dynamics of many tropical predators. Received: 28 June 1999 / Accepted: 29 February 2000     

Effects of egg size reduction and larval feeding on juvenile quality for a species with facultative-feeding development

In free-spawning marine invertebrates, larval development typically proceeds by one of two modes: planktotrophy (obligate larval feeding) from small eggs or lecithotrophy (obligate non-feeding) from relatively large eggs. In a rare third developmental mode, facultative planktotrophy, larvae can feed, but do not require particulate food to complete metamorphosis. Facultative planktotrophy is thought to be an intermediate condition that results from an evolutionary increase in energy content in the small eggs of a planktotrophic ancestor. We tested whether an experimental reduction in egg size is sufficient to restore obligate planktotrophy from facultative planktotrophy and whether the two sources of larval nutrition (feeding and energy in the egg) differentially influence larval survival and juvenile quality. We predicted, based on its large egg size, that a reduction in egg size in the echinoid echinoderm Clypeaster rosaceus would affect juvenile size but not time to metamorphosis. We reduced the effective size of whole (W) zygotes by separating blastomeres at the two- or four-cell stages to create half- (H) or quarter-size (Q) “zygotes” and reared larvae to metamorphosis, both with and without particulate food. Larvae metamorphosed at approximately the same time regardless of food or egg size treatment. In contrast, juveniles that developed from W zygotes were significantly larger, had higher organic content and had longer and more numerous spines than juveniles from H or Q zygotes. Larvae from W, H and Q zygotes were able to reach metamorphosis without feeding, suggesting that the evolution of facultative planktotrophy in C. rosaceus was accompanied by more than a simple increase in egg size. In addition, our results suggest that resources lost by halving egg size have a larger effect on larval survival and juvenile quality than those lost by withholding particulate food.     

Variation in maternal investment in a small cervid; the effects of cohort,sex, litter size and time of birth in roe deer (Capreolus capreolus) fawns

 We investigated the effects of cohort, sex, litter size and time of birth on birth weights and postnatal growth rates of roe deer fawns in a highly reproductive Norwegian population. By repeatedly recapturing radio-collared individuals, a total of 950 weights were obtained from 231 fawns of known age. In accordance with earlier studies, there was a period of linear growth during the first month following birth. Mean postnatal growth rates of 155 g/day are the highest yet recorded for roe deer; however, the mean birth weights of fawns were lower than those reported from populations in continental Europe. During the period of linear growth, we found no sex differences. However, growth rates were affected both by time of birth and litter size; fawns born early had lower growth rates than fawns born during or after the peak calving period, and fawns in triplet – groups had lower growth rates than either fawns in twin – groups or single fawns. Despite a fourfold increase in population density during the study, this factor was not able to explain variation in postnatal growth rates, although cohort effects on birth weight were evident. Received: 13 May 1996 / Accepted: 26 June 1996     

Growth and development of a generalist insect herbivore, Operophtera brumata, on original and alternative host plants

A generalist feeding strategy is common among eruptive insect herbivores but the ultimate reasons for a generalist strategy are not clear. Although generalist insect herbivores are able to complete their life cycle on several species of host plants, there is wide variation in the performance of individuals grown on different hosts. We examined whether different populations of Operophtera brumata are adapted to use the host species which is locally most abundant, and how the host plant affects growth and development of the insect. We reared two allopatric populations (eastern Finland, Prunus padus; south-west Finland, Quercus robur) on four species of host plants (Pr. padus, Populus tremula, Q. robur, Salix phylicifolia) from neonate larvae to the adult stage and measured the growth and development of individuals and the timing of adult hatching. The performance of both populations was best on Pr. padus, and the south-western population, originally on Q. robur, was well adapted to this host. The host affected the growth of females more than that of males. The host plant had an unexpected effect on hatching times of the adults. Individuals grown on the original host hatched in normal synchrony, i.e. males 6–7 days before females; but on alternative hosts this synchrony was disturbed. As is common in eruptive, capital-breeding generalist moths where female fecundity is linked to weight, host quality is critical for the flightless females of O. brumata. We suggest that in a heterogeneous environment the disturbing effect of alternative host plants on adult emergence may decrease the population density and growth rate compared to the potential maximum in a homogeneous environment. Received: 8 July 1999 / Accepted: 29 October 1999     

Effects of gestation temperature on offspring sex and maternal reproduction in a viviparous lizard (Eremias multiocellata) living at high altitude

Temperature-dependent sex determination (TSD) is well studied in many species of reptiles, but little is known on how geographic distribution and altitude affect the sex ratio. In the present study, we focused on a population of a viviparous lizard with TSD (Eremias multiocellata) that lives at high altitudes (≈2900 m) in Tianzhu, Gansu province, China. Gestation temperature had a notable effect on the offspring sex ratio, gestation period, and the mother's body mass. The mothers produced female biased offspring at 25 °C but male biased offspring at 35 °C. All female lizards lost weight during pregnancy, and the least loss of the body mass was observed at 31 °C. The gestation period increased in a non-linear fashion as ambient temperature was reduced. Average litter size was elevated with an increase of gestation temperatures, reached a maximum at 31 °C, and then declined at 35 °C. Compared with a previous study on a Minqin population which lives at a lower altitude (≈1400 m) and warmer climate, the present study obtained a less skewed sex ratio of offspring in the Tianzhu population. Geographic variations also affected offspring morphology between the two populations; females collected from Tianzhu produced larger litters but with a smaller body weight of offspring. These differences may be caused by the adaptive response to the cool climatic and high-altitude environmental conditions.     

Effects of salicylhydroxamic acid on the proliferation of Atriplex and murine neuroblastoma cells, and on Drosophila egg laying and development

Salicylhydroxamic acid (SHAM) inhibits the proliferation of cultured plant (Atriplex halimus) and murine neuroblastoma cells with IC50 of 90 and 250 microM, respectively. After 2 h of application, SHAM induces an acceleration of the neuroblastoma cell cycle from G1/S to G2 phases and, after 6 h, it induces an accumulation of the cells in S phase and a cell swelling. Up to 300 microM, SHAM is not cytotoxic and does not induce electrophysiological differentiation of neuroblastoma cells. When Drosophila females are grown in media containing 0.6-1.25 mM SHAM, the rate and number of laid eggs are increased. Furthermore, SHAM stimulates the different development stages from embryo to adult. A general interpretation of the effects of SHAM on cell proliferation and differentiation is proposed.     

The effect of embryonic development on metal and calcium content in eggs and eggshells in a small passerine

Published information relating to changes in the chemical element content of avian eggs caused by embryonic development is extremely scarce, although it may be crucial for understanding both the presence of anthropogenic pollutants as well as physiological levels of micronutrients. We assessed the variation in concentrations of calcium (Ca) and magnesium (Mg) and nine trace elements: seven essential (chromium (Cr), copper (Cu), nickel (Ni), manganese (Mn), iron (Fe), cobalt (Co) and zinc (Zn)) and two non‐essential (lead (Pb) and cadmium (Cd)) in shells and contents (both egg yolk and egg white) of embryonated and non‐embryonated eggs. We investigated the eggs of the Eurasian Reed Warbler Acrocephalus scirpaceus, a large proportion of whose eggs are infertile in our study population (almost 43% of clutches contain unhatched eggs) as well as significant embryo‐induced eggshell thinning at the equator of embryonated eggs. We found significantly higher concentrations (≥ 22.7%) of all the focal elements in the contents of embryonated eggs in comparison with non‐embryonated eggs, and a very pronounced one for Ca (nearly twice as high). The shells of embryonated eggs contained significantly higher concentrations of Zn (104.1%), Fe (56.5%), Pb (32.8%) and Cu (28.0%) but significantly lower ones of Co (8.9%) and Ca (9.3%) than the shells of non‐embryonated eggs. The simultaneous higher concentrations of all elements in the content of thinner‐shelled embryonated eggs suggest the parallel transfer of these elements along with Ca resorption from the shell into the egg interior during embryo formation. The higher concentration of most elements in the thinner shells of embryonated eggs may be indicative of the maternal deposition of some of these elements in a shell layer not subject to embryonic depletion, or in the eggshell membrane. Our results highlight the need for the careful selection of egg samples, which should differentiate between embryonated and non‐embryonated eggs in the analytical treatment of eggs and eggshells.