Publications

1993
Haig D. Alternatives to meiosis: the unusual genetics of red algae, microsporidia, and others. J Theor Biol. 1993;163 :15-31.Abstract

At a reductional division, the two alleles at a heterozygous locus segregate into different daughter nuclei. Reductional divisions are therefore vulnerable to genetic elements that cause one cell to attack its sister. This danger can be circumvented by creating uncertainty about when reduction occurs. In conventional meiosis, this is achieved by a sequence of two divisions with crossing over. Either the first or the second division can be reductional for most loci. In some red algae, reduction appears to be spread over more than two divisions. The novel reduction sequence of microsporidia creates maximal uncertainty about the time of reduction in two divisions without crossing over.

Haig D. Genetic conflicts in human pregnancy. Q Rev Biol. 1993;68 :495-532.Abstract

Pregnancy has commonly been viewed as a cooperative interaction between a mother and her fetus. The effects of natural selection on genes expressed in fetuses, however, may be opposed by the effects of natural selection on genes expressed in mothers. In this sense, a genetic conflict can be said to exist between maternal and fetal genes. Fetal genes will be selected to increase the transfer of nutrients to their fetus, and maternal genes will be selected to limit transfers in excess of some maternal optimum. Thus a process of evolutionary escalation is predicted in which fetal actions are opposed by maternal countermeasures. The phenomenon of genomic imprinting means that a similar conflict exists within fetal cells between genes that are expressed when maternally derived, and genes that are expressed when paternally derived. During implantation, fetally derived cells (trophoblast) invade the maternal endometrium and remodel the endometrial spiral arteries into low-resistance vessels that are unable to constrict. This invasion has three consequences. First, the fetus gains direct access to its mother's arterial blood. Therefore, a mother cannot reduce the nutrient content of blood reaching the placenta without reducing the nutrient supply to her own tissues. Second, the volume of blood reaching the placenta becomes largely independent of control by the local maternal vasculature. Third, the placenta is able to release hormones and other substances directly into the maternal circulation. Placental hormones, including human chorionic gonadotropin (hCG) and human placental lactogen (hPL), are predicted to manipulate maternal physiology for fetal benefit. For example, hPL is proposed to act on maternal prolactin receptors to increase maternal resistance to insulin. If unopposed, the effect of hPL would be to maintain higher blood glucose levels for longer periods after meals. This action, however, is countered by increased maternal production of insulin. Gestational diabetes develops if the mother is unable to mount an adequate response to fetal manipulation. Similarly, fetal genes are predicted to enhance the flow of maternal blood through the placenta by increasing maternal blood pressure. Preeclampsia can be interpreted as an attempt by a poorly nourished fetus to increase its supply of nutrients by increasing the resistance of its mother's peripheral circulation.

Haig D. Genomic imprinting, human chorionic gonadotropin, and triploidy. Prenat Diagn. 1993;13 :151.
1992
Haig D. Intragenomic conflict and the evolution of eusociality. J Theor Biol. 1992;156 :401-3.
1991
Haig D, Grafen A. Genetic scrambling as a defence against meiotic drive. J Theor Biol. 1991;153 :531-58.Abstract

Genetic recombination has important consequences, including the familiar rules of Mendelian genetics. Here we present a new argument for the evolutionary function of recombination based on the hypothesis that meiotic drive systems continually arise to threaten the fairness of meiosis. These drive systems act at the expense of the fitness of the organism as a whole for the benefit of the genes involved. We show that genes increasing crossing over are favoured, in the process of breaking up drive systems and reducing the fitness loss to organisms.

Haig D, Graham C. Genomic imprinting and the strange case of the insulin-like growth factor II receptor. Cell. 1991;64 :1045-6.
Moore T, Haig D. Genomic imprinting in mammalian development: a parental tug-of-war. Trends Genet. 1991;7 :45-9.Abstract

Genomic imprinting in mammals is increasingly being implicated in developmental and pathological processes, but without a clear understanding of its function in normal development. We believe that imprinting has evolved in mammals because of the conflicting interests of maternal and paternal genes in relation to the transfer of nutrients from the mother to her offspring. We present an hypothesis that accounts for many of the observed effects of imprinting in mammals and relates them to similar observations in plants. This hypothesis has implications for studies of X-chromosome inactivation and a range of human diseases.

Hurst LD, Haig D. Prion infection. Nature. 1991;351 :21.
Haig D, Hurst LD. A quantitative measure of error minimization in the genetic code. J Mol Evol. 1991;33 :412-7.Abstract

We have calculated the average effect of changing a codon by a single base for all possible single-base changes in the genetic code and for changes in the first, second, and third codon positions separately. Such values were calculated for an amino acid's polar requirement, hydropathy, molecular volume, and isoelectric point. For each attribute the average effect of single-base changes was also calculated for a large number of randomly generated codes that retained the same level of redundancy as the natural code. Amino acids whose codons differed by a single base in the first and third codon positions were very similar with respect to polar requirement and hydropathy. The major differences between amino acids were specified by the second codon position. Codons with U in the second position are hydrophobic, whereas most codons with A in the second position are hydrophilic. This accounts for the observation of complementary hydropathy. Single-base changes in the natural code had a smaller average effect on polar requirement than all but 0.02% of random codes. This result is most easily explained by selection to minimize deleterious effects of translation errors during the early evolution of the code.

1989
Haig D, Westoby M. Parent-specific gene expression and the triploid endosperm. Am Nat. 1989;134 :147-155.
Haig D, Westoby M. Selective forces in the emergence of the seed habit. Biol J Linn Soc. 1989;38 :215-238.
Haig D. Seed size and adaptation. Trends Ecol Evol. 1989;4 :145.
1988
Haig D, Westoby M. On limits to seed production. Am Nat. 1988;131 :757-759.
Haig D, Westoby M. A model for the origin of heterospory. J Theor Biol. 1988;134 :257-272.
Haig D, Westoby M. Sex expression in homosporous ferns: an evolutionary perspective. Evol Trends Plants. 1988;2 :111-119.
1987
Haig D. Kin conflict in seed plants. Trends Ecol Evol. 1987;2 :337-40.Abstract

Kin selection theory proposes that individuals value the reproductive success of relatives at a rate determined by their probability of shared alleles. The theory predicts when the interests of relatives are in accord and when they conflict. Though kin selection arguments have revolutionized the study of animal behavior, they have only recently been applied to plants. Kin selection has already been claimed to explain the formation of endosperm by double fertilization. This is the character that distinguishes angiosperms from gymnosperms. Plant life cycles involve interactions among kinds of relatives not encountered in animals. These interactions should be a fertile field for new applications of theory and the testing of ideas originally developed elsewhere.

1986
Haig D. Conflicts among megaspores. J Theor Biol. 1986;123 :471-480.

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