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De novo Genes in Plants: Origins, Mechanisms, and Functional Implications.

De novo genes originate from previously non-coding genomic regions. They provide an important source of lineage-specific innovation. In plants, these genes may contribute to adaptation, trait diversity and crop evolution. This review summarizes recent progress in plant de novo gene research. It first discusses major routes of gene birth, including transcription-first, open reading frame (ORF)-first and concurrent models. It also examines how nascent loci acquire regulatory control and enter existing biological networks. The review then summarizes their evolutionary features, including weak early constraint, rapid molecular change, restricted expression and structural refinement. It further discusses plant de novo genes involved in stress responses, seed germination, kernel dehydration, subspecies divergence, reproductive isolation and floral scent diversification. Current methods for identifying de novo genes remain limited by rapid sequence evolution, genome annotation quality, polyploidy and transposable elements. Whole-genome synteny alignment, multi-omics evidence and machine-learning approaches can improve candidate discovery. However, each method has important limitations. Finally, this review highlights key future questions in functional validation, latent coding potential in long non-coding RNAs, epigenetic activation, regulatory-network integration and crop improvement. These perspectives clarify how de novo genes shape plant adaptation and how they may be used in precision breeding and synthetic biology.

adaptive evolution

The Gibbons speciation mechanism.

A mechanism of sympatric speciation first proposed by Gibbons (1979, Am. Nat. 114, 719-741) is analyzed and submitted to computer simulation. It is found that, in its original form, the mechanism does not bring about reproductive isolation, but that with relatively minor modification, it may be made to work as claimed.

Animals

The inheritance of acquired epigenetic variations.

There is evidence that the functional history of a gene in one generation can influence its expression in the next. In somatic cells, changes in gene activity are frequently associated with changes in the pattern of methylation of the cytosines in DNA; these methylation patterns are stably inherited. Recent work suggests that information about patterns of methylation and other epigenetic states can also be transmitted from parents to offspring. This evidence is the basis of a model for the inheritance of acquired epigenetic variations. According to the model, an environmental stimulus can induce heritable chromatin modifications which are very specific and predictable, and might result in an adaptive response to the stimulus. This type of response probably has most significance for adaptive evolution in organisms such as fungi and plants, which lack distinct segregation of the soma and germ line. However, in all organisms, the accumulation of specific and random chromatin modifications in the germ line may be important in speciation, because these modifications could lead to reproductive isolation between populations. Heritable chromatin variations may also alter the frequency and distribution of classical mutations and meiotic recombination. Therefore, inherited epigenetic changes in the structure of chromatin can influence neo-Darwinian evolution as well as cause a type of "Lamarckian" inheritance.

Base Sequence

Allogeneic cellular reactions between intra-specific types of a solitary ascidian, Halocynthia roretzi.

Coelomic cells from a solitary ascidian, Halocynthia roretzi, exibit a nonphagocytic cellular reaction against coelomocytes from different species and another individuals of the same species. The reaction was denoted contact reaction. While xenogeneic contact reactions were always observed, allogeneic reactions were observed in most but not all combinations of individuals. Three variant types of H. roretzi (Type A, B, C) inhabit the coast of northern Japan. They are reproductively isolated under natural conditions. Non-reactive combinations exist between the different types, in all possible combinations. The paterns and frequencies of reactivities between different variants are almost the same to those observed within a single type. The results of the alloreactivity suggest that the three types of H. roretzi became separate from each other very recently and still remain intra-specific variants.

Alleles

Genic differentiation and origin of Robertsonian populations of the house mouse (Mus musculus domesticus Rutty).

This paper examines the relation between chromosomal and nuclear-gene divergence in 28 wild populations of the house mouse semi-species, Mus musculus domesticus, in Western Europe and North Africa. Besides describing the karyotypes of 15 of these populations and comparing them to those of 13 populations for which such information was already known, it reports the results of an electrophoretic survey of proteins encoded by 34 nuclear loci in all 28 populations. Karyotypic variation in this taxon involves only centric (or Robertsonian) fusions which often differ in arm combination and number between chromosomal races. The electrophoretic analysis showed that the amount of genic variation within Robertsonian (Rb) populations was similar to that for all-acrocentric populations, i.e. bearing the standard karyotype. Moreover, divergence between the two types of populations was extremely low. These results imply that centric fusions in mice have not modified either the level or the nature of genic variability. The genetic similarity between Rb and all-acrocentric populations is not attributed to the persistence of gene flow, since multiple fusions cause marked reproductive isolation. Rather, we attribute this extreme similarity to the very recent origin of chromosomal races in Europe. Furthermore, genic diversity measures suggest that geographically separated Rb populations have in situ and independent origins. Thus, Rb translocations are probably not unique events, but originated repeatedly. Two models are presented to explain how the rapid fixation of a series of chromosomal rearrangements can occur in a population without lowering variability in the nuclear genes. The first model assumes that chromosomal mutation rates are between 10(-3) and 10(-4) and that populations underwent a series of transient bottlenecks in which the effective population size did not fall below 35. In the second model, genic variability is restored following severe bottlenecks, through gene flow and recombination.

Animals

The divergence of a polygenic system subject to stabilizing selection, mutation and drift.

Polygenic variation can be maintained by a balance between mutation and stabilizing selection. When the alleles responsible for variation are rare, many classes of equilibria may be stable. The rate at which drift causes shifts between equilibria is investigated by integrating the gene frequency distribution W2N II (pq)4N mu-1. This integral can be found exactly, by numerical integration, or can be approximated by assuming that the full distribution of allele frequencies is approximately Gaussian. These methods are checked against simulations. Over a wide range of population sizes, drift will keep the population near an equilibrium which minimizes the genetic variance and the deviation from the selective optimum. Shifts between equilibria in this class occur at an appreciable rate if the product of population size and selection on each locus is small (Ns alpha 2 less than 10). The Gaussian approximation is accurate even when the underlying distribution is strongly skewed. Reproductive isolation evolves as populations shift to new combinations of alleles: however, this process is slow, approaching the neutral rate (approximately mu) in small populations.

Alleles

Mapping and characterization of a 'speciation gene' in Drosophila.

Almost nothing is known about the identity of the genes causing reproductive isolation between species. As a first step towards molecular isolation of a 'speciation gene', I mapped and partly characterized a gene causing hybrid male sterility in Drosophila. This analysis shows that sterility of D. melanogaster males who carry the 'dot' fourth chromosome from D. simulans is due entirely to a very small region of the D. simulans chromosome (including only about 5 salivary gland bands or approximately 250 kb of DNA). Thus the hybrid sterility effect of the D. simulans fourth chromosome is almost surely due to a single gene of very large effect (here named hms, hybrid male sterile). Hms is zygotically acting, and the D. simulans allele of hms is completely recessive. Furthermore, complementation tests suggest that hms is not an allele of any known locus in D. melanogaster.

Alleles

The genetic basis of Haldane's rule.

'Haldane's rule', formulated by J. B. S. Haldane in 1922, states that: "When in the F1 offspring of two different animal races one sex is absent, rare, or sterile, that sex is the heterozygous [heterogametic] sex". His rule is now known to apply in mammals, lepidopterans, birds, orthopterans and dipterans. In Drosophila, for example, Bock cites 142 cases of interspecific hybridizations that produce one sterile and one fertile sex in the offspring, all but one of these crosses yielding sterile XY males and fertile XX females. Despite much speculation, however, the genetic basis of Haldane's rule remains unknown. Haldane himself rejected the simple explanation that males are innately more sensitive than females to the effects of hybridization because groups with heterogametic females (such as birds and butterflies) usually show female sterility in hybrids, so that heterogamety itself is the critical feature. He and others suggested that heterogametic infertility or inviability in hybrids arises by a genetic imbalance between X chromosomes and autosomes. An alternative explanation is that this syndrome is caused by a mismatch of X and Y chromosomes. Here I show that in the Drosophila melanogaster subgroup, Haldane's rule for fertility apparently arises from a genetic interaction between X and Y chromosomes and not from an imbalance between sex chromosomes and autosomes. This finding has important implications for understanding the evolution of interspecific reproductive isolation.

Animals

Bidirectional incompatibility between conspecific populations of Drosophila simulans.

Cytoplasmic incompatibility (CI) describes the phenomenon whereby eggs fertilized by sperm from insects infected with a rickettsial endosymbiont fail to hatch. Unidirectional CI between conspecific populations of insects is a well documented phenomenon. Bidirectional CI has, however, only been described in mosquito populations, and recently between closely related species of parasitic wasps, where it is of interest as both an unusual form of reproductive isolation and as a potential means of insect population suppression. Here we report on the first known example of bidirectional CI between conspecific populations of Drosophila simulans. Further, we show that defects as early as the first cleavage division are associated with CI. This observation suggests that the cellular basis of CI involves disruption of processes before or during zygote formation and that CI arises from defects in the structure and/or function of the sperm during fertilization.

Animals

Location of an X-linked factor causing sterility in male hybrids of Drosophila simulans and D. mauritiana.

We report the first mapping of a genetic factor responsible for reproductive isolation: a small segment of genome strongly affecting sperm motility in hybrids between the sibling species Drosophila simulans and D. mauritiana. Maximum-likelihood analysis of data from ten generations of backcrossing places this factor at 1.1 +/- 0.2 map units from the forked locus, at position 54.9 +/- 0.2 or 57.1 +/- 0.2 on the X chromosome.

Animals

Genetic analysis of X-linked sterility in hybrids between three sibling species of Drosophila.

Three morphological markers (yellow, miniature, and forked) are used to map the location of X-chromosome segments causing male sterility in Drosophila simulans/D. mauritiana and D. simulans/D. sechellia hybrids. In both hybridizations at least three sections of the chromosome contain genes with substantial effects on sterility. This represents the maximum genetic divergence detectable with the three markers, suggesting that the X chromosome contains many loci affecting postzygotic reproductive isolation. The tight linkage between some markers and "sterility loci" may be useful in localizing and later cloning genes important in speciation.

Animals

Genetic variability of the interpulse interval of courtship song among some European populations of Drosophila melanogaster.

The interpulse interval of the courtship song of Drosophila melanogaster is a character which may play a significant role in mating success and reproductive isolation. Here we examine the variability of interpulse interval among replicated laboratory strains of D. melanogaster. There is no significant variation among populations of different geographical origin. This suggests that interpulse interval is subject to strong selection, as the populations are known to differ for other characters. One population, however, was sufficiently different to allow a genetic analysis. Reciprocal F1s and backcrosses implied that the variance was predominantly additive and autosomal. Possible sources of selection on interpulse interval are discussed.

Animals

[Original adaptive characters of intestinal Digenea of Sarpa salpa (Teleostei, Sparidae) and their interpretation in terms of evolution].

In the family Sparidae, the genus Sarpa is distinguished by a few characteristics: monospecificity, vegetarian diet and wide geographical distribution. The helminth fauna of Sarpa salpa is also very original. Indeed, the digenean parasites of this Teleostean fish are essentially classified into two families restricted to this fish. In the present paper, the author redescribes Mesometra orbicularis, M. brachycoelia, Centroderma spinosissima, Elstia stossichianum, Wardula capitellata (family Mesometridae) together with Robphildollfusium fractum (family Robphildollfusidae). Various original and yet unknown features are pointed out. Among these unusual structures, several correspond to adaptive characteristics favouring the settlement of the Digenean on the peculiar digestive gut wall of this herbivorous fish. Indeed, the intestinal mucous membrane of Sarpa salpa exhibits very few villi giving it an unusual smooth aspect. Therefore, the Mesometridae which always have just a single sucker (monostomatous) have selected a new kind of compensatory adhesive structure. Sometimes, the anterior end of the body becomes a sucker due to the particular distribution of the muscle strings; in other examples, the whole body becomes a sucker and its edges become considerably thinner to improve the tightness of the adhesive system. Other original anatomical features have been selected to allow survival in a medium rich in plant detritus. So, in the oral sucker crests ornemented by numerous sclerous denticles seem to act as a microfilter for the intestinal chyme in which plant fibres predominate. The original pharynx seems to act as a suction-force pump. The excretory system, which is of a reticular type, penetrates the whole parenchyma and this could be a response to huge intestinal fermentations. The Digenea of Sarpa salpa are not interpreted by the author as true parasites but as endocommensal symbionts. These inquiline species are not immunogenic, or at least only slightly so, since they do not feed upon the host itself but upon its intestinal chyme. In most cases this results in a high parasite density (post larvae and adults) together with a cohabitation of the various species along the various intestinal segments. Coexistence of several species, systematically very close, evidently raises the question of their reproductive isolation. The author proposes an answer founded upon data of allopatric speciation.

Adaptation, Physiological

Pole cells of Drosophila paulistorum: embryologic differentiation with symbionts.

The pole cells of young D. paulistorum embryos are destined to form the germinal cells of both male and female imagoes. In addition, specialized portions of the midgut may be derived from pole cell progenitors. In this initial study of their embryogenesis by means of electron microscopy, various stages of pole cell development are shown in both non-hybrid (potentially fertile) and intersemispecific hybrid (potentially sterile as males) materials. Originally, approximately 5 or 6 cells emerge to form the early polar cap and subsequently divide asynchronously until the 35-50 cells of the late polar cap are derived. Unlike other Drosophila species, however, mycoplasma-like symbionts, apparently an hereditary infection, have been traced to locations within the cytoplasm of these pole cells. They are depicted as arriving there after transmission via the egg cytoplasm, implicating this as their probable route of entry into the future germinal tissues of adult flies. It is postulated that these microorganisms function as an infectious reproductive isolating mechanism fostering hybrid male sterility between D. paulistorum semispecies.

Animals

Interlocus variation of genetic distance and the neutral mutation theory.

Theoretical distributions of genetic distance between reproductively isolated taxa are derived by means of computer simulation, taking into account mutation and random genetic drift. The distributions obtained are in good agreement with the observed distributions of interracial and interspecific genetic distances for enzymes loci in Drosophila. This indicates that the gene substitution at enzyme loci can be explained by the neutral mutation theory.

Animals

Differing levels of dispersed repetitive DNA among closely related species of Drosophila.

The genomic concentrations of certain middle repetitive DNA sequences vary considerably among closely related species of Drosophila. In fact, the chromosomes of D. melanogaster appear to carry approximately 3 times as much middle repetitive DNA as those of the sibling species D. simulans. Although most of the middle repetitive DNA of D. melanogaster consists of segments of "nomadic" DNA that occupy different dispersed chromosomal locations in different strains of flies, repeated DNA sequences recovered from the D. simulans genome are most often restricted to single chromosomal positions. Apparent differences in the total concentrations of middle repetitive DNA in the two species are most easily attributed to an approximately sevenfold difference in their dispersed repetitive and nomadic DNA contents. These differences may affect the relative mutation rates of these species or contribute to their reproductive isolation or both.

Animals

On the association of restriction fragment length polymorphisms across species boundaries.

We study the expected values of gametic-phase disequilibrium in both nuclear and cytonuclear systems in a finite population composed of reproductively isolated subpopulations. Random drift alone within each subpopulation will generate permanent overall non-zero gametic-phase disequilibrium in both a two-locus nuclear system and a cytonuclear system unless the population has an overall initial disequilibrium of zero. We derive formulae for the expected overall disequilibrium when mutation is involved and demonstrate that these values decay to zero at an extremely slow rate compared with the decay of the expected disequilibria within each subpopulation.

Animals

Ancient and recent patterns of geographic speciation in the oyster mushroom Pleurotus revealed by phylogenetic analysis of ribosomal DNA sequences.

Evidence from molecular systematic studies suggests that many mushroom species may be quite ancient. Gene phylogenies were developed to examine the relationship between reproductive isolation, genetic divergence, and biogeography in oyster mushrooms (Pleurotus). Sequence data were obtained for two regions of DNA from populations belonging to eight intersterility groups (biological species). Phylogenetic analysis of sequences from the 5' portion of the nuclear encoded large subunit rDNA demonstrates an ancient origin for four intersterility groups of broad geographic distribution (world-wide), with a more recent radiation of several intersterility groups that are restricted to the Northern Hemisphere. An expanded analysis using sequence data from the more variable rDNA internal transcribed spacer region also reveals a phylogenetically based pattern of genetic divergence associated with allopatric speciation among populations from different continents in the Northern Hemisphere. The ability of rDNA sequences to resolve phylogenetic relationships among geographically isolated populations within intersterility groups illustrates the importance of biogeography for understanding speciation in Pleurotus. Patterns of geographic distribution among intersterility groups suggest that several species lineages evolved quite early, with recently evolved groups restricted to the Northern Hemisphere and older lineages occurring throughout the world. Based on phylogenetic evidence, analysis of historical biogeography using area cladograms shows that multiple dispersal and vicariance events are responsible for patterns of speciation observed.

Base Sequence