Effects of delayed fertilization on embryo viability in walleye Sander vitreus (Mitchill): the role of maternal effects

The effects of delayed fertilization on embryonic viability in walleye Sander vitreus were examined using laboratory fertilization and incubation trials. Both fertilization success (survival to 10 h post-fertilization) and embryonic survival (from 10 to 168 h post-fertilization) generally declined with increasing ova storage time but trends varied significantly among females. Embryonic survival of individual egg batches was positively related to fertilization success. The slope of the relationship between fertilization success and ova storage time was negatively related to maternal fork length indicating that ova quality declined more rapidly with storage time for larger females than for smaller females. In contrast, the slope of the early embryonic survival v. ova storage time relationship was not significantly related to maternal size.     

Inbreeding, outbreeding and environmental effects on genetic diversity in 46 walleye (Sander vitreus) populations

Genetic diversity is recognized as an important population attribute for both conservation and evolutionary purposes; however, the functional relationships between the environment, genetic diversity, and fitness-related traits are poorly understood. We examined relationships between selected lake parameters and population genetic diversity measures in 46 walleye (Sander vitreus) populations across the province of Ontario, Canada, and then tested for relationships between six life history traits (in three categories: growth, reproductive investment, and mortality) that are closely related to fitness, and genetic diversity measures (heterozygosity, d2, and Wright's inbreeding coefficient). Positive relationships were observed between lake surface area, growing degree days, number of species, and hatchery supplementation versus genetic diversity. Walleye early growth rate was the only life history trait significantly correlated with population heterozygosity in both males and females. The relationship between FIS and male early growth rate was negative and significant (P < 0.01) and marginally nonsignificant for females (P = 0.06), consistent with inbreeding depression effects. Only one significant relationship was observed for d2: female early growth rate (P < 0.05). Stepwise regression models showed that surface area and heterozygosity had a significant effect on female early growth rate, while hatchery supplementation, surface area and heterozygosity had a significant effect on male early growth rate. The strong relationship between lake parameters, such as surface area, and hatchery supplementation, versus genetic diversity suggests inbreeding and outbreeding in some of the populations; however, the weak relationships between genetic diversity and life history traits indicate that inbreeding and outbreeding depression are not yet seriously impacting Ontario walleye populations.     

Interaction of sauger Sander canadensis and walleye Sander vitreus in a large,shallow northern river

The objectives of this study were to: (1) determine the relative abundance of sauger Sander canadensis and walleye S. vitreus within the Rainy River using a standardized index netting technique; (2) assess the life-history characteristics of the two species in a northern river; (3) examine the spatial distribution of the two species; (4) assess year-class synchrony. At a larger scale, relative abundance of S. canadensis and S. vitreus were similar among sections of the river. However, at a finer scale, S. canadensis were the dominant species over more area (24%) of the river than S. vitreus (12%). Life-history traits for both species were within the range reported for North American. Mortality rates were similar, suggesting that anglers were not affecting one species more than the other. Year-class strengths were not synchronous between S. canadensis and S. vitreus (r = 0.07). There was evidence of S. canadensis potentially outcompeting S. vitreus with a strong year class of S. canadensis followed by a very weak class of S. vitreus. Additionally, S. canadensis were larger than S. vitreus in most sections of the river when adjusted for mean size. However, the potential interspecific competition was not to the exclusion of S. vitreus. Turbidity was probably the factor that enable S. canadensis to survive sympatrically with S. vitreus given their inability to segregate by depth within the river.     

Exposure to low concentrations of dissolved ammonia promotes growth rate in walleye Sander vitreus

The objective of the current study was to examine whether sublethal (moderate) levels of dissolved ammonia may be beneficial to growth in juvenile walleye Sander vitreus (recent evidence in juvenile rainbow trout Oncorhynchus mykiss has shown significant increases in protein synthesis in the presence of moderately elevated concentrations of dissolved ammonia). Moderately elevated dissolved ammonia concentrations between 100 and 300 μmol l⁻¹ suppressed routine aerobic metabolic activity by 20% during acute trials (2 h), while promoting specific growth rate (>50%) and elevating whole body soluble protein content by 20% in the early stages (14–42 days) in chronic ammonia exposure experiments. Juvenile S. vitreus held at ammonia concentrations between 107·6 ± 5·5 and 225·5 ± 4·7 μmol l⁻¹ (mean ± s . e .) grew significantly faster than control fish and significantly reduced plasma cortisol levels (<3 μg dl⁻¹). Results from this study suggest that chronic exposure to moderate amounts of dissolved ammonia significantly increase growth rates in juvenile S. vitreus by increasing nitrogen accessible for supplementary protein deposition leading to somatic development.     

Characterization and expression of a myosin heavy–chain isoform in juvenile walleye Sander vitreus

In this study, myosin, the major component of myofibrillar protein in the skeletal muscle, was characterized and its expression was monitored during growth in juvenile walleye Sander vitreus. First, the coding region of myosin heavy chain (MyHC) from the fast skeletal muscle of walleye was amplified by long-distance PCR using a full-length cDNA. Phylogenetic analysis was used to determine the evolutionary relationship of this S. vitreus myosin sequence to other vertebrate myosin sequences. Next, it was established that the myosin isoform was most prevalent in the white muscle, compared with the red and cardiac muscle. Myosin expression was monitored over a series of experiments designed to influence growth. Specifically, change in MyHC mRNA was monitored after acute changes in feeding. Fish exposed to a one-week fasting period showed significant decreases in MyHC mRNA levels by the end of the fast. The effect of feeding was also examined more closely over a 24 h period after feeding, but results showed no significant change in myosin expression levels through this time period. Finally, fish with higher growth rates had higher MyHC mRNA and protein expression levels. This study indicates that MyHC mRNA expression is sensitive to the factors that may influence growth in juvenile S. vitreus .     

Temporal effective size estimates of a managed walleye Sander vitreus population and implications for genetic-based management

The goal of this research was to use the long-term fishery data set and DNA from archived scales of walleye Sander vitreus in Escanaba Lake, WI, U.S.A., to improve the understanding of the underlying mechanism(s) influencing genetic diversity in naturally recruiting populations. The introduced population of S. vitreus in Escanaba Lake has a low mean effective population size ( N _E) between 124·6 and 185·5 despite a mean census size ( N _C) of 4659 ( N _E/ N _C c. 0·04), suggesting an accelerated rate of genetic drift between 1952 and 2002. These values are smaller than the median N _E range of several studies suggesting typical N _E/ N _C ratios of 0·11–0·16 in a wide range of taxa. N _E increased steadily during the past two sampled decades (1992 and 2002) and was consistent with a lowering of the variance in S. vitreus reproductive success, possibly linked to a large, sustained exploitation (mean 28%) rate. Variance in reproductive success is one of the most important factors influencing N _E in species, like S. vitreus , which have a potential for large fecundities and large juvenile mortalities (type III survivorship). The N _B estimates across six sequential cohorts (age classes of S. vitreus , assayed from 1994 to 1999) was consistent with estimates of N _E reported for 1992–2002. These results, coupled with in-depth census and exploitation data, show that the genetic characteristics of Escanaba Lake S. vitreus have changed substantially and that management activities, such as supplemental stocking and harvest practices, have profoundly influenced the genetic dynamics of S. vitreus in this lake.     

Maternal condition influences phenotypic selection on offspring

1. Environmentally induced maternal effects are known to affect offspring phenotype, and as a result, the dynamics and evolution of populations across a wide range of taxa. 2. In a field experiment, we manipulated maternal condition by altering food availability, a key factor influencing maternal energy allocation to offspring. We then examined how maternal condition at the time of gametogenesis affects the relationships among early life-history traits and survivorship during early development of the coral reef fish Pomacentrus amboinensis. 3. Maternal condition did not affect the number of embryos that hatched or the number of hatchlings surviving to a set time. 4. We found no significant difference in egg size in relation to the maternal physiological state. However, eggs spawned by supplemented mothers were provisioned with greater energy reserves (yolk-sac and oil globule size) than nonsupplemented counterparts, suggesting that provision of energy reserves rather than egg size more closely reflected the maternal environment. 5. Among offspring originating from supplemented mothers, those with larger yolk-sacs were more likely to successfully hatch and survive for longer periods after hatching. However, among offspring from nonsupplemented mothers, yolk-sac size was either inconsequential to survival or offspring with smaller yolk-sac sizes were favoured. Mothers appear to influence the physiological capacity of their progeny and in turn the efficiency of individual offspring to utilize endogenous reserves. 6. In summary, our results show that the maternal environment influences the relationship between offspring characteristics and survival and suggest that energy-driven selective mechanisms may operate to determine progeny viability.     

Signatures of vicariance, postglacial dispersal and spawning philopatry: population genetics of the walleye Sander vitreus

Population genetic relationships reveal the signatures of current processes such as reproductive behaviour and migration, as well as historic events including vicariance and climate change. We analyse population structure of native walleye Sander vitreus across North America, encompassing 10 nuclear DNA microsatellite loci, 26 spawning sites and 921 samples from watersheds across the Great Lakes, Lake Winnipeg, upper Mississippi River, Ohio River and Mobile Bay of the Gulf Coast. Geographical patterning is assessed using phylogenetic trees, pairwise F _ST analogues, hierarchical partitioning, Mantel regression, Bayesian assignment and Monmonier geographical networks. Results reveal congruent divergences among population groups, corresponding to historic isolation in glacial refugia, dispersal patterns and basin divisions. Broad-scale relationships show genetic isolation with geographical distance, but reproductive groups within basins do not – with some having pronounced differences. Greatest divergence distinguishes outlying Gulf Coastal and northwest populations, the latter tracing to dispersal from the Missourian refugium to former glacial Lake Agassiz, and basin isolation ∼7000 ya. Genetic barriers in the Great Lakes separate groups in Lakes Superior, Huron's Georgian Bay, Erie and Ontario, reflecting contributions from Mississippian and Atlantic refugia, and changes in connectivity patterns. Walleye genetic patterns thus reflect vicariance among watersheds and glacial refugia, followed by re-colonization pathways and changing drainage connections that established modern-day northern populations, whose separations are maintained through spawning site fidelity. Conservation management practices should preserve genetic identity and unique characters among these divergent walleye populations.     

Maternal body condition influences magnitude of anti-predator response in offspring

Organisms exhibit plasticity in response to their environment, but there is large variation even within populations in the expression and magnitude of response. Maternal influence alters offspring survival through size advantages in growth and development. However, the relationship between maternal influence and variation in plasticity in response to predation risk is unknown. We hypothesized that variation in the magnitude of plastic responses between families is at least partly due to maternal provisioning and examined the relationship between maternal condition, egg provisioning and magnitude of plastic response to perceived predation risk (by dragonfly larvae: Aeshna spp.) in northern leopard frogs (Lithobates pipiens). Females in better body condition tended to lay more (clutch size) larger (egg diameter) eggs. Tadpoles responded to predation risk by increasing relative tail depth (morphology) and decreasing activity (behaviour). We found a positive relationship between morphological effect size and maternal condition, but no relationship between behavioural effect size and maternal condition. These novel findings suggest that limitations imposed by maternal condition can constrain phenotypic variation, ultimately influencing the capacity of populations to respond to environmental change.     

Toxicological effects of methylmercury on walleye (Sander vitreus) and perch (Perca flavescens) from lakes of the boreal forest

Biochemical and physiological responses of walleye (Sander vitreus) and perch (Perca flavescens) were studied in four Canadian boreal forest lakes representing a mercury (Hg) exposure gradient. The aim of this study was to assess the effects of Hg and methylmercury (MeHg) on the general physiological condition of fish as well as to gauge the relationship between MeHg and the glutathione (GSH) system in metal-contaminated and reference sites using a series of biomarkers. Walleye from Lake Malartic had the highest liver MeHg concentrations, exhibited lower hepatosomatic indices (HSI) and lower glutathione S-transferase (GST) activity. HSI was negatively related to liver total Hg concentrations in walleye (R2=0.33, n=108, P<0.0001). Glutathione reductase (GR) and GST activity for walleye from Lake Malartic were related to HSI (R2=0.38, n=25, P=0.0010; R2=0.46, n=27, P<0.0001, respectively). In Lake Desjardins-East, where perch had the highest liver MeHg concentrations, glutathione peroxidase selenium dependent activity (GSH-Px SD) and GST activity were negatively related to liver MeHg concentrations (R2=0.39, n=21, P=0.0026; R2=0.22, n=21, P=0.0298, respectively). This study suggests that Hg may induce adverse effects on the physiology and cellular metabolism of walleye and perch at environmentally relevant concentrations.     

Interplay between ecological, behavioural and historical factors in shaping the genetic structure of sympatric walleye populations (Sander vitreus)

Disentangling ecological, behavioural and evolutionary factors responsible for the presence of stable population structure within wild populations has long been challenging to population geneticists. This study primarily aimed at decoding population structure of wild walleye (Sander vitreus) populations of Mistassini Lake (Québec, Canada) in order to define source populations to be used for the study of spatial partitioning using individual-based multilocus assignment methods, and decipher the dynamics of individual dispersal and resulting patterns of spatial resource partitioning and connectivity among populations. A second objective was to elucidate the relationships between biological characteristics (sex, size, age and population of origin) and an individual's probability to migrate and/or disperse. To do so, a total of 780 spawning individuals caught on five distinct spawning sites, and 1165 postspawning individuals, captured over two sampling seasons (2002-2003) were analysed by means of eight microsatellite loci. Four temporally stable walleye populations associated with distinct reproductive grounds were detected. These populations were differentially distributed among lake sectors during their feeding migration and their spatial distribution was stable over the two sampling seasons. Dispersing individuals were identified (n=61); these revealed asymmetrical patterns of dispersal between populations, which was also confirmed by divergent admixture proportions. Regression models underlined population of origin as the only factor explaining differential dispersal of individuals among populations. An analysis of covariance (ancova) indicated that larger individuals tended to migrate from their river of origin further away in the lake relative to smaller fish. In summary, this study underlined the relevance of using individual-based assignment methods for deciphering dynamics of connectivity among wild populations, especially regarding behavioural mechanisms such as differential spatial partitioning and dispersal responsible for the maintenance of genetic population structure.     

Purification and properties of Sandercyanin, a blue protein secreted in the mucus of blue forms of walleye, Sander vitreus

A blue protein present in the mucus coating blue forms of walleye, Sander vitreus, was purified to homogeneity by a combination of ion exchange and hydrophobic interaction chromatography. The purified protein has a molecular mass of 87,850 and is a homotetramer with a subunit molecular mass of 21,836. Solutions of the protein are deep blue in color and show absorbance maxima at 383 and 633 nm, respectively. Acetone treatment of the protein releases a blue chromophore with the spectral characteristics of biliverdin IXα. The results suggest that the blue protein is a new biliprotein that we have termed Sandercyanin. The function of Sandercyanin is unknown, but limited amino acid sequence analysis suggests it is a lipocalin that may be involved in the transport of heme degradation products.

Maternal allocation strategies and differential effects of yolk carotenoids on the phenotype and viability of yellow-legged gull (Larus michahellis) chicks in relation to sex and laying order

Egg quality may mediate maternal allocation strategies according to progeny sex. In vertebrates, carotenoids have important physiological roles during embryonic and post-natal life, but the consequences of variation in yolk carotenoids for offspring phenotype in oviparous species are largely unknown. In yellow-legged gulls, yolk carotenoids did not vary with embryo sex in combination with egg laying date, order and mass. Yolk lutein supplementation enhanced the growth of sons from first eggs but depressed that of sons from last eggs, enhanced survival of daughters late in the season, and promoted immunity of male chicks and chicks from small eggs. Lack of variation in egg carotenoids in relation to sex and egg features, and the contrasting effects of lutein on sons and daughters, do not support the hypothesis of optimal sex-related egg carotenoid allocation. Carotenoids transferred to the eggs may rather result from a trade-off between opposing effects on sons or daughters.     

Optimization of fertilization success in Sander vitreus is influenced by the sperm : egg ratio and ova storage

Walleye Sander vitreum eggs could be stored at 7° C for up to 30 h with 60% fertilization rates. Fertilization rates at 1 and 12° C fell to 17 and 5%, respectively. At spermatozoa : egg ratios <25 000 : 1, survival to the eyed stage declined precipitously. Beyond a ratio of 25 000 : 1, survival to the eyed stage did not increase > c . ∼ 70%. Therefore, a standard spermatozoa : egg ratio of 25 000 : 1 was established and is recommended for studies concerning fertilizing success.     

Age at maturity in wild baboons: genetic, environmental and demographic influences

The timing of early life-history events, such as sexual maturation and first reproduction, can greatly influence variation in individual fitness. In this study, we analysed possible sources of variation underlying different measures of age at social and physical maturation in wild baboons in the Amboseli basin, Kenya. The Amboseli baboons are a natural population primarily comprised of yellow baboons (Papio cynocephalus) that occasionally hybridize with anubis baboons (Papio anubis) from outside the basin. We found that males and females differed in the extent to which various factors influenced their maturation. Surprisingly, we found that male maturation was most strongly related to the proportion of anubis ancestry revealed by their microsatellite genotypes: hybrid males matured earlier than yellow males. In contrast, although hybrid females reached menarche slightly earlier than yellow females, maternal rank and the presence of maternal relatives had the largest effects on female maturation, followed by more modest effects of group size and rainfall. Our results indicate that a complex combination of demographic, genetic, environmental, and maternal effects contribute to variation in the timing of these life-history milestones.     

A Study of Maternal and Maternal Interaction Effects in Diallel Crosses

The analysis of diallel crosses by including the components due to maternal effect and maternal interaction effects have been presented for Griffing method—1 (random effect model) and Griffing method—3 (fixed and random effect model). Wherever exact test of significance is not possible, testing procedure using Satterthwaite (1946) approximation has been presented.     

The effect of mycorrhizal infection of Abutilon theophrasti on competitiveness of offspring

1. Our goal was to determine whether mycorrhizal infection of one generation of plants could influence the competitiveness of the subsequent generation.
2. We grew the offspring of mycorrhizal (M) and non-mycorrhizal (NM) Abutilon theophrasti plants together in dense populations in large boxes in a greenhouse.
3. Offspring of M plants were larger than offspring of NM plants. As the experiment progressed, the difference in size was magnified and self-thinning occurred.
4. Compared with offspring of NM plants, offspring of M plants had a twofold greater survival and collectively produced a total of nearly four times as many seeds.
5. We conclude that mycorrhizal infection of the parent generation can increase the competitive ability of the offspring.     

Maternal and embryonic sources of tyrosine hydroxylase during Drosophila embryogenesis

Tyrosine hydroxylase (TH), the enzyme which catalyzes the conversion of tyrosine to L-DOPA and is rate limiting in catecholamine biosynthesis, is biochemically expressed in late stage wild-type Drosophila oocytes as well as in early embryogenesis. Null mutant alleles of TH (pale) are embryonic lethals with death occurring in the late embryonic or early larval periods of development. Staging of embryos demonstrated that inhibition of the enzymatic activity of TH by alpha-methyl-p-tyrosine (alphaMT) retards the progression of embryos primarily during the organogenesis stages of embryonic development, with lesser effects on earlier and later stages. On the other hand, time of gene action studies with a conditional temperature sensitive pale mutant (ple(ts1)) at its restrictive temperature (29 degrees C) indicate an onset of tyrosine hydroxylase gene action beginning in the oocyte stage of development. Thus, maternal as well as embryonic effects on the secretion and/or functionality of this enzyme may play roles in the early developmental program of the organism.     

Evolution of sex-biased maternal effects in birds. IV. Intra-ovarian growth dynamics can link sex determination and sex-specific acquisition of resources

The evolutionary importance of maternal effects is determined by the interplay of maternal adaptations and strategies, offspring susceptibility to these strategies, and the similarity of selection pressures between the two generations. Interaction among these components, especially in species where males and females differ in the costs and requirements of growth, limits inference about the evolution of maternal strategies from their expression in the offspring phenotype alone. As an alternative approach, we examine divergence in the proximate mechanisms underlying maternal effects across three house finch populations with contrasting patterns of sex allocation: an ancestral population that shows no sex-biased ovulation, and two recently established populations at the northern and southern boundaries of the species range that have opposite sequences of ovulation of male and female eggs. For each population, we examined how oocyte acquisition of hormones, carotenoids and vitamins was affected by oocyte growth and overlap with the same and opposite sexes. Our results suggest that sex-specific acquisition of maternal resources and sex determination of oocytes are linked in this system. We report that acquisition of testosterone by oocytes that become males was not related to growth duration, but instead covaried with temporal exposure to steroids and overlap with other male oocytes. In female oocytes, testosterone acquisition increased with the duration of growth and overlap with male oocytes, but decreased with overlap with female oocytes. By contrast, acquisition of carotenoids and vitamins was mostly determined by organism-wide partitioning among oocytes and oocyte-specific patterns of testosterone accumulation, and these effects did not differ between the sexes. These results provide important insights into three unresolved phenomena in the evolution of maternal effects - (i) the evolution of sex-specific maternal allocation in species with simultaneously developing neonates of both sexes; (ii) the link between sex determination and sex-specific acquisition of maternal products; and (iii) the evolution of context-dependent modulation of maternal effects.