Kinship and covariance

Price's (1970) covariance theorem can be used to derive an expression for gene frequency change in kin selection models in which the fitness effect of an act is independent of the genotype of the recipient. This expression defines a coefficient of relatedness which subsumes r(Wright, 1922), b(Hamilton, 1972), ρ (Orlove &; Wood, 1978), and R(Michod &; Hamilton, 1980). The new coefficient extends the domain of Hamilton's rule to models in which the average gene frequency of actors differs from that of recipients.     

Helping in cooperatively breeding long-tailed tits: a test of Hamilton's rule

Inclusive fitness theory provides the conceptual framework for our current understanding of social evolution, and empirical studies suggest that kin selection is a critical process in the evolution of animal sociality. A key prediction of inclusive fitness theory is that altruistic behaviour evolves when the costs incurred by an altruist (c) are outweighed by the benefit to the recipient (b), weighted by the relatedness of altruist to recipient (r), i.e. Hamilton''s rule rb > c. Despite its central importance in social evolution theory, there have been relatively few empirical tests of Hamilton''s rule, and hardly any among cooperatively breeding vertebrates, leading some authors to question its utility. Here, we use data from a long-term study of cooperatively breeding long-tailed tits Aegithalos caudatus to examine whether helping behaviour satisfies Hamilton''s condition for the evolution of altruism. We show that helpers are altruistic because they incur survival costs through the provision of alloparental care for offspring. However, they also accrue substantial benefits through increased survival of related breeders and offspring, and despite the low average relatedness of helpers to recipients, these benefits of helping outweigh the costs incurred. We conclude that Hamilton''s rule for the evolution of altruistic helping behaviour is satisfied in this species.     

The genetic mating structure of subdivided populations I. Open-mating model

The genetic mating structure of a subdivided population can describe how parental genotypes gave rise to zygotes. When parents of the same genotype are considered together as one class (“open-mating”), three independent parameters of inbreeding and mating structure are needed to describe this structure at a diallelic locus. One is Wright's fixation index F. The other two are mating structure parameters, derived herein and termed the “effective selfing” rate E and the “inbreeding assortative selfing” rate D. E is the genetically equivalent proportion of self-fertilization at a single locus, and is given by standardized second and third central moments of gene frequencies of mates. E is a summary measure of inbreeding that includes effects due to self-fertilization and mating to relatives, as well as correlations between mates induced by Wahlund effects and/or selective diversification among neighborhoods. The second parameter D measures the tendency of inbred or more homozygous individuals to effectively self more (or less) than outbred or more heterozygous individuals. D is related to the maintenance of variation of inbreeding among individuals and/or to the prevalence of spatial variation of selection. D is independent of E, but together with E controls the generational change of inbreeding, ΔF. Extensions of the model to unequal allele frequencies in male vs female mates, and to multi-allelic loci, are also examined.     

The "cytochrome b5 fold": structure of a novel protein superfamily

Selective proteolysis allows the isolation of a heme-binding fragment spectrally similar to microsomal cytochrome b₅ from both baker's yeast flavocytochrome b₂ (a flavohemoprotein) and liver sulfite oxidase (a molybdoprotein). The amino acid sequences of these two fragments have been published separately (Guiard &; Lederer, 1976,1979). We present in this paper an alignment of those sequences with that of microsomal cytochrome b₅. The structural consequences of the similarity between the three primary structures are discussed in the light of the cytochrome b₅ three-dimensional model (Mathews et al., 1971,1972,1975; Mathews &; Czerwinski, 1976).It is concluded that the three heme-binding proteins are in all probability the products of a divergent evolution from a common ancestor and that they must present a basically similar backbone with some surface alterations. We propose to name this backbone the “cytochrome b₅ fold”. The comparison of the three proteins suggests hypotheses concerning the molecular surface areas involved in the recognition of cytochrome c (the common acceptor) and of the respective reductase (flavo- or molybdoprotein).In addition, our results suggest that at some point in evolution, several copies of an initial hemoprotein gene were formed in the cellular genome. Subsequently, one copy was fused with the gene for another function: a flavoreductase in yeast cells or a molybdoreductase in hepatic cells.     

An elementary treatment of the genetical theory of kin-selection

Hamilton's (coefficient of relation) rule for the spread of a gene influencing altruistic behavior is derived under a simple model (discrete generations, single locus, infinite population, panmictic mating) using elementary probability theory. The rule holds quite well with slow selection and it makes no difference whether the gene is dominant or recessive. The rule is altered, however, if the pool of relatives (in interaction) is finite, and I derive a correction factor for interaction between sibs in a sexual haploid. In general the ratio of benefit to cost (necessary for the altruist gene to be favored) increases as the family size decreases.     

A theoretical study of calcium entry in nerve terminals, with application to neurotransmitter release

In his study of a kin selection model for the evolution of workers' behavior in an incipiently social insect, Craig (1979) found that the haplodiploid mode of sex determination, combined with a female-biased sex ratio, cannot accelerate an evolutionary trend toward eusociality. This seems to contradict to Hamilton's (1964) theory. It is my intention to prove that Craig's result is due to the dependence of relative reproductive success in each sex on the sex ratio of a population. The oviposition of unfertilized eggs by workers is indispensable, in primitive stages in the evolution of eusociality, for maintaining the relative reproductive success of a female. Assuming that workers control the sex ratios, we can distinguish the following three evolutionary phases in accordance with the ratio $\text{N}{}_2\text{N}{}_{10}$ (the ratio of the number of the queen's second brood to that of the progeny laid by workers derived from the same nest). During Phase 1 in which $\text{N}{}_2\text{N}{}_{10}$ < $\text{1}\text{2}$, the reproductive successes of a male and a female are equal, and Hamilton's rule holds. In Phase 2 in which $\text{1}\text{2 <}\text{N}{}_2\text{N}{}_{10}\text{<}\text{3}\text{2}$, the queen produces females, and workers oviposit males. As $\text{N}{}_2\text{N}{}_{10}$ increases, the sex ratio in the whole population becomes female-biased and relative reproductive success of a male increases. In Phase 3 in which N₂/N₁₀ >$\text{3}\text{2}$, the queen lays some male eggs in addition to female eggs. The sociality threshold ($\text{B}\text{C}$)_crit the minimum value of the benefit/cost ratio leading to the evolution of altruism, is $\text{2}\text{3}$on Phase 1. It rises threefold as N₂/N₁₀ increases in Phase 2, and, in Phase 3, it is twice as high as that in a diploid species. The anatomical and physiological characteristics of workers must have been so developed that they are efficient in helping activities after the beginning of eusociality. The evolutionary process before the beginning of helping behavior is also discussed. The egg laying by unmated females, which stay their mother's nest, seems to have been an important preadaptation for the evolution of eusociality in Hymenoptera.     

Male‐biased gene flow across an avian hybrid zone: evidence from mitochondrial and microsatellite DNA

Mating pattern and gene flow were studied in the contact zone between two morphologically very similar Chiffchaff taxa (Phylloscopus collybita, P. brehmii) in SW France and northern Spain. Mating was assortative in brehmii, but not in collybita. Mixed matings were strongly asymmetric (excess of collybita male × brehmii female pairs), but did produce viable offspring in some cases. Sequence divergence of the mitochondrial cytochrome b gene was 4.6%. Haplotypes segregated significantly with phenotype (only five ‘mismatches’ among 94 individuals), demonstrating that mitochondrial gene flow was very restricted. The estimated proportion of F₁ hybrids in the reproductive population was significantly lower than expected under a closed population model, indicating strong selection against hybrids. Genetic typing of 101 individuals at four microsatellite loci also showed significant population differentiation, but nuclear gene flow was estimated to be 75 times higher than mitochondrial gene flow. This strong discrepancy is probably due to unisexual hybrid sterility (Haldane’s rule). Thus, there is a strong, but incomplete, reproductive barrier between these taxa.     

Gene mutations in rye causing embryo lethality in hybrids with wheat: allelism and chromosomal localisation

In crosses between hexaploid wheat and inbred lines of rye, a small number of rye genotypes produce seeds carrying undifferentiated, non-viable embryos. Hybrids between such lines and those not showing this phenotype were used as pollen donors in crosses with bread wheat in order to determine the genetic basis of disturbed embryo development. A single gene, designated Eml-R1b, is causing this character. Molecular markers associated with F₂ genotypes derived from a contrasting rye inbred progeny were used for a linkage study. Recombinant inbred lines of an F₅ population served as testers. Eml-R1b maps to chromosome arm 6RL, along with two co-segregating microsatellite loci, Xgwm1103 and Xgwm732. Complementary interactions of deleterious genes in wheat and rye are discussed.     

Genetic mapping and QTL analysis for yield and agronomic traits with an F2:3 population derived from a waxy corn × sweet corn cross

We constructed a framework map using SSR markers in the F₂ population derived from a cross between a waxy corn inbred line and a sweet corn inbred line. We constructed a genetic linkage map of the F_2:3 population employing 295 SSR markers on 158 F₂ individuals produced from the cross. The map comprised a total genomic length of 2,626.5 cM in 10 linkage groups and an average distance between markers of 8.9 cM. The number of loci per linkage group ranged from 27 (chr. 5) to 34 (chr. 7). The genetic distance per linkage group ranged from 213.6 cM (chr. 10) to 360.6 cM (chr. 2). Χ ² tests revealed that 254 markers (86.1 %) distributed over all 10 chromosomes exhibited a Mendelian segregation ratio of 1:2:1. A total of 14 quantitative trait loci (QTLs) for days to silking (DTS), plant height (PH), ear height (EH), ear height ratio (ER), ear length (L-ear), and setted ear length (L-sear) were found in the 158 F₂ progeny. They were mapped to chromosomes 1, 2, 3, 7, 8, and 10. Among them, one QTL was associated with DTS, three with PH, six with EH, one with ER, two with L-ear, and one QTL was related to L-sear. In our study, we found that four QTLs: qDTS1, qEH1a, qEH1b, and qPH1, were clustered between umc2390 and umc1603 on chromosome 1. These new QTLs identified by the present study could serve as useful molecular markers in selecting for yield and agronomic traits in maize. The results of this study may improve the identification and characterization of genes responsible for yield and agronomic traits in waxy corn and sweet corn.     

Evolution of social behavior by reciprocation

A general model is presented for the evolution of social behavior by reciprocation. The results of our model apply to social traits which are transmitted from one generation to the next by a process which guarantees that the frequency of the trait in one generation is directly related to its fitness in the preceding generation. The basic parameters of the model are α, the number of interactions per generation, and β, the number of these interactions which are with individuals who are perceived as strangers. It is shown that so long as α/β can be made large, social reciprocation may increase when arbitrarily rare even in the absence of population structure.This conclusion appears to be at odds with several recent investigations of Axelrod & Hamilton (1981) and Boorman & Levitt (1980). We use our model to reconcile these various approaches. By casting Axelrod & Hamilton's (1981) single-partner model in terms of the general parameters, α and β, we show that social reciprocation can increase when arbitrarily rare in a homogeneous population dominated by non-cooperators. Using a gene frequency approach, Boorman & Levitt (1973, 1980) demonstrated the existence of a selection threshold in frequency of the social trait, which must be surmounted for social reciprocation to increase. We show our analysis of reciprocation to be consistent with Boorman and Levitt's result, since for our general model the cost to the social individuals of learning the non-social's identity goes to zero as the ratio α/β gets large. We also use our general model to study two multi-partner models not considered elsewhere, which differ in regards to the memory capabilities assigned to the organism.Finally we use our model to compare directly the evolution of social behavior by reciprocation with the main alternate hypothesis, kin selection. We show that an act which accrues some cost ?c to the fitness of the donor while benefiting a recipient by b, will increase in frequency so long as c/b < Φ (equation (30)), where Φ is defined as the “coefficient of reciprocation” or probability that a cooperative act is reciprocated. By comparing the coefficient of reciprocation with the coefficient of relatedness of kin selection, direct comparisons of the two hypotheses may be made.     

淮河野生鲇鱼线粒体Cyt b基因的序列变异与遗传结构分析

本研究利用线粒体DNA细胞色素b(Cyt b)基因序列分析淮河信阳段、淮滨段、蚌埠段、洪泽湖及其支流颍河、淠河和池河的野生鲇鱼(Silurus asotus)种群遗传结构及种群历史.结果表明,在841 bp的同源序列中,7个种群共检测到变异位点40个,占全部序列的4.76％,121个个体共检测到32种单倍型;7个种群的平均单倍型多样性(h)、核甘酸多样性(Pi)分别为0.884 8、0.003 8,表明淮河野生鲇鱼种群的遗传多样性水平较高.7个种群间的遗传分化指数Fst为0.115 0,仅12.92％的变异来自种群间(AMOVA分析),基因交流值为3.85,种群间K2-P遗传距离为0.002 ～0.009,显示鲇鱼种群间没有发生明显的地理分化.NJ树揭示7个种群的个体组成2个谱系,但这2个谱系与地理分布并不相关.中性检验、错配分析和Network网络亲缘关系分析皆表明鲇鱼种群有过种群扩张,扩张时间约在0.17～0.29百万年前的中更新世中期和末期.     

Studies on the orientations of the mitochondrial redox carriers. Orientation of the chromophores of cytochrome b--c1 complex with respect to the plane of a cytochrome b--c1 complex--lipid model membrane.

Orientations of the active site chromophores of the mitochondrial cytochrome b-c₁ complex incorporated into liposomes have been investigated in hydrated oriented multilayers of proteoliposome membranes using optical and epr spectroscopy. The hemes of cytochromes c₁ and b were found to be oriented with the normal to their heme planes lying approximately in the plane of the proteoliposome membrane. Rieske's iron-sulfur center was oriented with the z-axis of the g tensor parallel to the plane of the membranes. It is concluded that the cytochrome b-c₁ complex has a structural asymmetry which causes it to orient with respect to the lipid bilayer.     

Body temperature and basking behaviour of Nile crocodiles (Crocodylus niloticus) during winter

The ability to thermoregulate in reptilians is often through behavioural modification. We investigated body temperature (T_b) patterns during winter in the amphibious Nile crocodile (Crocodylus niloticus) and its relationship to basking behaviour at the St. Lucia Crocodile Centre, St. Lucia, South Africa. It was found that crocodiles had no daily plateaus in T_b but rather continuous oscillations in T_b within a range of mean minimum T_b 18.8–19.6 °C to mean maximum T_b 26.9–29.2 °C. Crocodile T_b increased during the day, usually after 10:00 irrespective of body size. Behavioural data showed that the crocodiles usually left the water to bask around 10:00. It is suggested that basking behaviour is important for elevating T_b rather than attaining a preferred T_b. The increased T_b may allow them to perform optimally when they return to water. The basking occurrence has management implications as it suggests that the best time to conduct aerial censuses of the St. Lucia crocodiles is during winter after 10:00 when most of the individuals are basking and hence most easily seen.     

The effect of multiple inseminations on the evolution of social behaviors in diploid and haplo-diploid organisms

Using a classical population genetic model, the necessary conditions for the spread of genes that determine social behaviors and the rate of spread of these genes are derived. The influence of 1, 2, 3, or k inseminations per female on these conditions is investigated for both diploid and haplodiploid organisms. These results are then extended to a population in which there are arbitrary variations among females in their numbers of mates. These results do not depend upon assuming equal paternity by all inseminating males; the effects of sperm competition and unequal paternity are also derived. The rates and conditions for social evolution in these groups of complex composition are discussed in relation to Hamilton's rule.For all models, the total change in gene frequency, Δq, is partitioned into two components: (1) $\text{Δq}{}_{\mathrm{I}}$, the change in gene frequency caused by selection within groups; this component is always negative, illustrating that individual selection always operates against the evolution of social behaviors; and (2) Δq_G, the change in gene frequency caused by selection between groups; this component is generally positive. Hamilton's rule is shown to specify the necessary and sufficient conditions for $\text{Δq}{}_{\mathrm{G}}\text{> |}\text{Δq}{}_{\mathrm{I}}\text{|}$, that is, for selection among kin groups to over-ride individual selection within kin groups.     

DNA,chromatin and chromosomes: By E. M. Bradbury,N. Maclean and H. R. Matthews. New York: Halsted Press (a division of John Wiley and Sons). (1981). 281 pp. $24.95

The Ah locus regulates the induction of cytochrome P₁-450 by foreign chemicals such as 3-methyl-cholanthrene and 2,3,7,8-tetrachlorodibenzo-p-dioxin. The induction process is controlled by the cytosolic Ah receptor. The cytosolic and nuclear Ah receptors were studied in the liver from inbred C57BL/6N (Ah^b/Ah^b) mice, inbred DBA/2N (Ah^d/Ah^d) mice and heterozygotes (Ah^b/Ah^d) and homozygotes (Ah^d/Ah^d) derived from the (C57BL/6N × DBA/2N)F1 × DBA/2N backcross. After [³H-1,6]-2,3,7,8-tetrachlorodibenzo-p-dioxin (³H-TCDD) is given in vivo, the receptor in Ah^b/Ah^b and Ah^b/Ah^d mice is detectable in the cytoplasm and nucleus; in Ah^d/Ah^d mice the receptor is not measurable in the cytosol, but is found in the nucleus at levels one fourth to one fifth of those in Ah^b/Ah^b mice. P₁-450 (23S) mRNA content was estimated by Northern hybridization and by Rot analysis with a mouse P₁-450 cloned cDNA. An excellent dose-response relationship (r = 0.99) was found between the amount of ³H-TCDD-Ah receptor complex appearing in the nucleus and the quantity of P₁-450 mRNA induced in mice with all three possible Ah genotypes.     

Inclusive fitness and the sociobiology of the genome

Inclusive fitness theory provides conditions for the evolutionary success of a gene. These conditions ensure that the gene is selfish in the sense of Dawkins (The selfish gene, Oxford University Press, Oxford, 1976): genes do not and cannot sacrifice their own fitness on behalf of the reproductive population. Therefore, while natural selection explains the appearance of design in the living world (Dawkins in The blind watchmaker: why the evidence of evolution reveals a universe without design, W. W. Norton, New York, 1996), inclusive fitness theory does not explain how. Indeed, Hamilton’s rule is equally compatible with the evolutionary success of prosocial altruistic genes and antisocial predatory genes, whereas only the former, which account for the appearance of design, predominate in successful organisms. Inclusive fitness theory, however, permits a formulation of the central problem of sociobiology in a particularly poignant form: how do interactions among loci induce utterly selfish genes to collaborate, or to predispose their carriers to collaborate, in promoting the fitness of their carriers? Inclusive fitness theory, because it abstracts from synergistic interactions among loci, does not answer this question. Fitness-enhancing collaboration among loci in the genome of a reproductive population requires suppressing alleles that decrease, and promoting alleles that increase the fitness of its carriers. Suppression and promotion are effected by regulatory networks of genes, each of which is itself utterly selfish. This implies that genes, and a fortiori individuals in a social species, do not maximize inclusive fitness but rather interact strategically in complex ways. It is the task of sociobiology to model these complex interactions.     

Intermembrane electron transport in the absence of added water-soluble carriers

Electron transport from untreated to mersalyzed microsomal vesicles at the level of NADH-cytochrome b₅ reductase or cytochrome b₅ has been demonstrated in the absence of added water-soluble electron carriers. A similar effect was shown in the systems “intact mitochondria — mersalyzed microsomes” and “mersalyzed mitochondria— untreated microsomes”. No measurable electron transport between intact and mersalyzed particles of inner mitochondrial membrane was found. The obtained data suggest that the capability to carry out intermembrane electron transfer is specific for NADH-cytochrome b₅ reductase and/or cytochrome b₅, localized in microsomal and outer mitochondrial membranes.     

Spatial Genetic Structure of Campanula sabatia, a Threatened Narrow Endemic Species of the Mediterranean Basin

Campanula sabatia is endemic to NW Italy (Liguria) and it was included in the European Red list of endangered species due to the heavy human pressure on its habitat. AFLP markers were used to detect the genetic diversity within and among ten populations (totalling 83 individuals) representing the range of the species. In spite of the limited distribution of this endemic taxon, high levels in percentage of polymorphic bands (PPB), gene diversity (H _S and H _T) and Shannon??s information index (I) were detected both at population (PPB?=?60?%, H _S?=?0.1853, I?=?0.2836) and at species level (PPB?=?100?%, H _T?=?0.2415, I?=?0.3871). The coefficient of genetic differentiation among population (G??_ST) was 0.1935, while the level of gene flow (N?? _m) was estimated to be 2.0832. AMOVA analysis identified a genetic variation within populations of 83?% of the total. Bayesian clustering methods assigned individuals to two geographical groups partly found within the same population, probably due to a high rate of genetic exchange among its populations. Conservation measures are suggested on the basis of the genetic diversity detected to ensure an effective protection for this endemic species.     

Thermal preferences of hatchling saltwater crocodiles (Crocodylus porosus) in response to time of day,social aggregation and feeding

Three month old hatchling Crocodylus porosus with data loggers in their stomachs were placed in thermal gradients, in isolation (N=16) and in groups of 4 (N=8 groups; 32 individuals). Mean T_b and variation in T_b (SD) was not different whether individual crocodiles in isolation were fasted or fed, or if individuals were housed in isolation (I) or in groups (G). However, individuals in isolation (N=16) maintained slightly lower T_bs than those in groups (N=32) during the early morning (06:00–11:00 h). The overall mean T_b recorded for fasted individuals in the isolated and group treatments (N=48) was 30.9±2.3 °C SD, with 50% of T_bs (T_set) between 29.4 °C and 32.6 °C, and a voluntary maximum and minimum of 37.6 °C and 23.2 °C respectively. During the day (11:00–17:00 h), individuals in isolation and in groups selected the warmer parts of the gradient on land, where they moved little. Outside of this quiescent period (QP), activity levels were much higher and they used the water more. There was a strong diurnal cycle for fasted individuals in isolation and in groups, with T_b during the QP (31.9±2.09 °C; N=48) significantly higher than during the non-quiescent period (NQP: 30.6±2.31 °C). Thermal variation (SD) in T_b was relatively stable throughout the day, with the highest variation at around dusk and early evening (18:00–20:00 h), which coincided with a period of highest activity. The diurnal activity cycle appears innate, and may reflect the need to engage in feeding activity at the water's edge in the early evening, despite ambient temperatures being cooler, with reduced activity and basking during the day. If so, preferred T_b may be more accurately defined as the mean T_b during the QP rather than the NQP. Implications for the thermal environment best suited for captive C. porosus hatchlings are discussed.