The evolution of biogeochemistry: revisited

Biogeochemistry - The evolution of biogeochemistry, retraces the important historical steps in part, covered by Gorham (Biogeochemistry 13:199–239, 1991) in the 18–19th...     

Ancient asymmetries in the evolution of flowers

Dorsoventral asymmetry in flowers is thought to have evolved many times independently as a specialized adaptation to animal pollinators. To understand how such a complex trait could have arisen repeatedly, we have compared the expression of a gene controlling dorsoventral asymmetry in Antirrhinum with its counterpart in Arabidopsis, a distantly related species with radially symmetrical flowers. We found that the Arabidopsis gene is expressed asymmetrically in floral meristems, even though they are destined to form symmetrical flowers. This suggests that, although the flowers of the common ancestor were probably radially symmetrical, they may have had an incipient asymmetry, evident at the level of early gene activity, which could have been recruited many times during evolution to generate asymmetric flowers.     

MADS about the evolution of orchid flowers

Orchids have unique flowers involving three types of perianth organs: outer tepals, lateral inner tepals, and a lip. Expression studies indicate that the identity of these organs is specified by the combinatorial interaction of four different DEFICIENS-like MADS-box genes. We suggest that clarifying the evolution of these genes provides a rational framework for reconstructing the enigmatic origin and unique diversification of the orchid flower. For example, two rounds of gene duplications during early orchid evolution might have generated the genes that were probably recruited to distinguish the different types of perianth organs. This hypothesis suggests intriguing, experimentally testable mechanisms by which gene duplications followed by sub- and neo-functionalization events might have contributed to the evolutionary origin of morphological novelties in orchids - and well beyond.     

The evolution of dominance: Haldane v Fisher revisited.

Fisher's model for the evolution of dominance indicates that the accumulation of dominance modifiers will be accelerated by (1) an increased frequency of the mutant heterozygote, (2) increased selection for the phenotype of the normal homozygote. The model has been criticised by Haldane on the grounds that point (1) is not fulfilled, that is dominance appears to be more common in populations with a low frequency of mutant heterozygotes (populations of inbreeders). In support of Fisher's model it is argued that intense selection for the wild type phenotype is more common in inbreeders than outbreeders. This situation should promote the accumulation of dominance modifiers (point (2) above).     

The evolution of altruism between siblings: Hamilton's rule revisited

This paper explores the validity of Hamilton's rule in the case of other-only altruism in which the benefits are shared by other members of the sibling group excluding the donor. It presents a model of competition between two alleles which code for different kinds of altruism. It derives a simple replicator equation for allele frequencies under conditions of strong selection. This equation does not depend on the size of the sibling group. In mathematical form, the equation is similar to Hamilton's original rule in the case of inbreeding, although the causal mechanism is different. The paper derives a simple criterion to determine whether there will be a polymorphism in which both alleles coexist permanently. Such an event is rare and victory will normally go to the allele with the higher value of 1/2b-c, where b is the total benefit which an offspring confers on its siblings and c is the cost to the donor. The paper also considers how an offspring will behave in particular circumstances. Using a specialized version of the basic model, it shows how, in the absence of polymorphism, natural selection should take the system towards the point of 50% marginal altruism. With this type of altruism, an offspring will perform any act for which the expected cost to the donor is at most half the expected benefit to its siblings. Acts which do not satisfy this criterion are not performed. This accords with Haldane's quip that he would sacrifice his own life for two of his brothers, but not for less. Numerical simulation is used to explore these issues in greater depth. The paper also examines briefly the implications of heterozygote advantage for Hamilton's rule. It concludes with a brief discussion of the connection between other-only altruism and whole-group altruism, in which the donor gains some benefit from its actions.     

Buzz-pollination in three nectariferousBoraginaceae and possible evolution of buzz-pollinated flowers

Buzz-pollination was observed in three nectariferousBoraginaceae spp.:Onosma gigantea Lam.,Trichodesma africana (L.)R. Br. andT. boissieri Post. An evolutionary pathway from usual nectariferous flowers to typical buzz-pollinated flowers is suggested.     

Heterochrony revisited: the evolution of developmental sequences

The concept of heterochrony is a persistent component of discussions about the way that evolution and development interact. Since the late 1970s heterochrony has been defined largely as developmental changes in the relationship of size and shape. This approach to heterochrony, here termed growth heterochrony, is limited in the way it can analyse change in the relative timing of developmental events in a number of respects. In particular, analytical techniques do not readily allow the study of changes in developmental events not characterized by size and shape parameters, or of many kinds of events in many taxa. I discuss here an alternative approach to heterochrony, termed sequence heterochrony, in which a developmental trajectory is conceptualized as a series of discrete events. Heterochrony is demonstrated when the sequence position of an event changes relative to other events in that sequence. I summarize several analytical techniques that allow the investigation of sequence heterochrony in phylogenetic contexts and also quantitatively. Finally, several examples of how this approach may be used to test hypotheses on the way development evolves are summarized.     

The evolution of unisexual flowers: morphological and functional convergence results from diverse developmental transitions

Unisexual flower morphology was examined within a phylogenetic context in order to identify developmental transitions associated with the multiple origins of dioecy in flowering plants. Historically, two categories of unisexual flowers have been recognized: type I flowers exhibit rudiments of the nonfunctional organ type, while type II flowers bear no vestigial sexual organs. Mapping of these flower types onto a composite phylogeny shows that type II morphology is homoplasious and has resulted from at least four distinct evolutionary developmental pathways. The historical assignment of unisexual flowers into only two morphological types has masked important developmental and evolutionary dynamics.     

Animal evolution: the enigmatic phylum placozoa revisited

A recent report of high levels of genetic variation between strains of Trichoplax adhaerens challenges the traditional view that the phylum Placozoa comprises only one species. At the morphological level, placozoans are amongst the simplest extant animals, but molecular evidence suggests that they may have more complex origins.     

The evolution of life histories in spatially heterogeneous environments: Optimal reaction norms revisited

Summary Natural populations live in heterogeneous environments, where habitat variation drives the evolution of phenotypic plasticity. The key feature of population structure addressed in this paper is the net flow of individuals from source (good) to sink (poor) habitats. These movements make it necessary to calculate fitness across the full range of habitats encountered by the population, rather than independently for each habitat. As a consequence, the optimal phenotype in a given habitat not only depends on conditions there but is linked to the performance of individuals in other habitats. We generalize the Euler-Lotka equation to define fitness in a spatially heterogeneous environment in which individuals disperse among habitats as newborn and then stay in a given habitat for life. In this case, maximizing fitness (the rate of increase over all habitats) is equivalent to maximizing the reproductive value of newborn in each habitat but not to maximizing the rate of increase that would result if individuals in each habitat were an isolated population. The new equation can be used to find optimal reaction norms for life history traits, and examples are calculated for age at maturity and clutch size. In contrast to previous results, the optimal reaction norm differs from the line connecting local adaptations of isolated populations each living in only one habitat. Selection pressure is higher in good and frequent habitats than in poor and rare ones. A formula for the relative importance of these two factors allows predictions of the habitat in which the genetic variance about the optimal reaction norm should be smallest.     

Accelerated rates of floral evolution at the upper size limit for flowers

Evolutionary theory explains phenotypic change as the result of natural selection, with constraint limiting the direction, magnitude, and rate of response [1]. Constraint is particularly likely to govern evolutionary change when a trait is at perceived upper or lower limits. Macroevolutionary rates of floral-size change are unknown for any angiosperm family, but it is predicted that rates should be diminished near the upper size limit of flowers, as has been shown for mammal body mass [2]. Our molecular results show that rates of floral-size evolution have been extremely rapid in the endoholoparasite Rafflesia, which contains the world's largest flowers [3]. These data provide the first estimates of macroevolutionary rates of floral-size change and indicate that in this lineage, floral diameter increased by an average of 20 cm (and up to 90 cm)/million years. In contrast to our expectations, it appears that the magnitude and rate of floral-size increase is greater for lineages with larger flowered ancestors. This study suggests that constraints on rates of floral-size evolution may not be limiting in Rafflesia, reinforcing results of artificial- and natural-selection studies in other plants that demonstrated the potential for rapid size changes [4-6].     

The genetics of mirror-image flowers

Conspicuous asymmetries in forms that are polymorphic within a species can be genetically or environmentally determined. Here, we present a genetic analysis of the inheritance of dimorphic enantiostyly, a sexual polymorphism in which all flowers on a plant have styles that are consistently deflected either to the left or the right side of the floral axis. Using Heteranthera multiflora (Pontederiaceae), a short-lived herb, we conducted crosses within and between left- and right-styled plants and scored progeny ratios of the style morphs in F(1), F(2) and F(3) generations. Crosses conducted in the parental generation between morphs or right-styled plants resulted in right-styled progeny, whereas crosses between left-styled plants resulted in left-styled progeny. When putative heterozygous F(1) plants were selfed, the resulting F(2) segregation ratios were not significantly different from a 3 : 1 ratio for right- and left-styled plants. Crosses between left- and right-styled plants in the F(2) generation yielded F(3) progeny with either a 1 : 1 ratio of left- and right-styled plants or right-styled progeny. Our results are consistent with a model in which a single Mendelian locus with two alleles, with the right-styled allele (R) dominant to the left-styled allele (r), governs stylar deflection. The simple inheritance of dimorphic enantiostyly has implications for the evolution and maintenance of this unusual sexual polymorphism.     

The evolution and loss of oil-offering flowers: new insights from dated phylogenies for angiosperms and bees

The interactions between bees that depend on floral oil for their larvae and flowers that offer oil involve an intricate mix of obligate and facultative mutualisms. Using recent phylogenies, new data on oil-offering Cucurbitaceae, and molecular-dating, we ask when and how often oil-offering flowers and oil-foraging bees evolved, and how frequently these traits were lost in the cause of evolution. Local phylogenies and an angiosperm-wide tree show that oil flowers evolved at least 28 times and that floral oil was lost at least 36–40 times. The oldest oil flower systems evolved shortly after the K/T boundary independently in American Malpighiaceae, tropical African Cucurbitaceae and Laurasian Lysimachia (Myrsinaceae); the ages of the South African oil flower/oil bee systems are less clear. Youngest oil flower clades include Calceolaria (Calceolariaceae), Iridaceae, Krameria (Krameriaceae) and numerous Orchidaceae, many just a few million years old. In bees, oil foraging evolved minimally seven times and dates back to at least 56 Ma (Ctenoplectra) and 53 Ma (Macropis). The co-occurrence of older and younger oil-offering clades in three of the four geographical regions (but not the Holarctic) implies that oil-foraging bees acquired additional oil hosts over evolutionary time. Such niche-broadening probably started with exploratory visits to flowers resembling oil hosts in scent or colour, as suggested by several cases of Muellerian or Batesian mimicry involving oil flowers.     

On evolution of andromonoecy and 'overproduction' of flowers: a resource allocation model

The conditions for the evolution of andromonoecy and male-function-controlled overproduction of fertile flowers in hermaphrodites are considered using the evolutionarily stable strategy (ESS) approach. Andromonoecy and male-function-controlled floral display are promoted if further increase in pollen amount per flower is disadvantageous and/or increase in the number of polliniferous units is advantageous; the costs of attractive organs and pollen per flower are relatively low while the cost of ovules per flower and the cost per fruit are relatively high; the probability of setting a fruit from a pistillate flower is high; male fertility increases with the resources devoted to flowering; and if selfing is relatively low. Assumptions and predictions of the model are discussed.     

The evolution of diadromy in fishes (revisited) and its place in phylogenetic analysis

Diadromy is a term used to describe migrations of fishes between fresh waters and the sea; these migrations are regular, physiologically mediated movements which occur at predictable life history phases in each diadromous species, they involve most members of a species' populations, and they are usually obligatory. Around 250 fish species are regarded as diadromous. A review of the life history strategies amongst families of fishes that include diadromous species provides little support for a suggested scenario for their evolution that involves: (1) evolution of anadromy via amphidromy from fishes of marine origins, and (2) evolution of catadromy through amphidromy from fishes of freshwater origins, even though these scenarios seem intuitively reasonable. The various forms of diadromy appear to have had multiple independent origins amongst diverse fish groups. There is increasing confidence that behavioural characteristics of animals are heuristic in gener ating and interpreting phylogenies. However, examination of fishes shows wide variability of diadromous life histories within closely related families and genera, within species, and there is even ontogenetic variation in patterns of behaviour by individual fish. In addition, there is multiple loss of diadromy in many diadromous fish species in which the life history becomes restricted to fresh waters. This variation suggests that diadromy is a behavioural character of dubious worth in determining phylogenetic relationships. Moreover, it appears to have been an ancestral condition in some fish families, such as Anguillidae, Salmonidae, Galaxiidae, Osmeridae, and others, and perhaps in the whole salmonoid/osmeroid/galaxioid complex of families. This, too, makes diadromy of dubious worth in phylogenetic analysis