The S locus of flowering plants: when self-rejection is self-interest.

In certain families of flowering plants, a self-incompatibility (SI) locus prevents self-fertilization, by a specific interaction between the S-gene product produced in the pistil and the S-gene products borne on or expressed by the male gametophyte, the pollen grain. The female S-locus gene products for two families showing different types of SI have been putatively identified as major pistil glycoproteins (the S-locus-specific glycoproteins of the Brassicaceae and the S-RNases of the Solanaceae). However, they are distinct in sequence and mode of action. The nature of the S-locus gene product borne by the pollen is still uncertain in both systems.     

Self-fertilization and the evolution of recombination

In this article, we study the effect of self-fertilization on the evolution of a modifier allele that alters the recombination rate between two selected loci. We consider two different life cycles: under gametophytic selfing, a given proportion of fertilizations involves gametes produced by the same haploid individual, while under sporophytic selfing, a proportion of fertilizations involves gametes produced by the same diploid individual. Under both life cycles, we derive approximations for the change in frequency of the recombination modifier when selection is weak relative to recombination, so that the population reaches a state of quasi-linkage equilibrium. We find that gametophytic selfing increases the range of epistasis under which increased recombination is favored; however, this effect is substantial only for high selfing rates. Moreover, gametophytic selfing affects the relative influence of different components of epistasis (additive x additive, additive x dominance, dominance x dominance) on the evolution of the modifier. Sporophytic selfing has much stronger effects: even a small selfing rate greatly increases the parameter range under which recombination is favored, when there is negative dominance x dominance epistasis. This effect is due to the fact that selfing generates a correlation in homozygosity at linked loci, which is reduced by recombination.     

Cytological Characterization of Self Incompatibility in Gametes of the Ascidian, Ciona intestinalis

We have determined how many elements are involved in the regulation of self-fertilization in the solitary ascidian, Ciona intestinalis that is an incompletely self-sterile species. Animals collected in the field were repeatedly induced to spawn in order to examine their selfing ratios. About 20% of them were self-fertile, although the ratios fluctuated considerably among respective spawnings. Naturally or acid-induced self-fertile gametes required much longer time for selfing than that for crossing. Egg-suspending seawater (egg water) as such activated sperm motility, but it lowered conspicuously the self-fertilization ratio. Self-sterile spermatozoa could scarcely bind to the vitelline coat (VC) of glycerinated autologous eggs. or in case they bound well to it the sperm flagella ceased to beat within five min of the 'insemination'. The staining of sectioned gametes with DAPI, a fluorescent dye of DNA, showed that in selfing the spermatozoa could hardly penetrate the VC even though they bound well to it. The results of this study show that the block of self-fertilization can be classified into four elements from a phenomenal viewpoint, such as egg water, low affinity of sperm-VC binding, inactivation of bound sperm and difficulty in sperm penetration through the VC.     

Inheritance of parental genomes by a hybrid form <Emphasis Type="Italic">Rana “esculenta”</Emphasis> (Amphibia,Ranidae)

In this study, quantitative analysis of paternal genome inheritance by a hybrid form Rana “esculenta” (= Rana esculenta L., 1758 × Rana ridibunda Pall., 1881) (Amphibia, Ranidae) was examined. The hybrid form examined was characterized by a polymodal mode of inheritance (genome of any of the parental species can be inherited). The absence of correlation between the proportion of normal gametes and either sex or ploidity of the producer was demonstrated. The gametes produced could be both haploid and diploid (hybrid or homozygous). The mechanism of alloploid reproduction is discussed.     

Additive by Additive Variance with Inbreeding and Linkage

Two-locus coancestries, which provide the coefficients of the additive by additive component in the variance and covariance of relatives for a quantitative trait, were formulated generally in terms of two-locus probabilities of identity by descent for combinations of parental and recombinant gametes. Explicit expressions, with linkage, were developed for all relatives of self-fertilization, for selfed and outbred relatives and for full and half sibs from unrelated inbred parents. The relative effect of linkage on the two-locus coancestry decreases rapidly as inbreeding and relatedness of the relatives increase. It was concluded that the error caused by ignoring linkage would probably be small in the estimation of genetic variances or in the projection of selection response with self-fertilization.     

Complex patterns of mating revealed in a Eucalyptus regnans seed orchard using allozyme markers and the neighbourhood model

The neighbourhood model apportions offspring of individual mother plants to self-fertilization, outcrossing to males within a circumscribed area around the mother plant (the neighbourhood), and outcrossing to males outside the neighbourhood. Formerly the model was applied only to haploid pollen gametes in the offspring of conifers, but is extended so that it can be used with genotypic data from diploid offspring of both angiosperms and gymnosperms. In addition, it is shown that the mating parameters can be estimated without independent estimates of allele frequencies in the pollen pools outside the neighbourhood; thus the model might be applied effectively to natural populations exposed to unknown external pollen sources. Parameters of the neighbourhood mating model were estimated for a 10-year-old seed orchard population of the insect-pollinated tree, Eucalyptus regnans, in southeast Australia, which contained a mixture of two geographical provenances (Victoria and Tasmania). The mating patterns revealed were complex. Crosses between trees of the same provenance occurred three times more often than crosses between trees of different provenances. Levels of self-fertilization and patterns of mating within neighbourhoods were influenced by provenance origin, crop fecundity and orchard position (central vs. edge) of mother trees. Gene dispersal, however, was extensive, with approximately 50% of effective pollen gametes coming from males more than 40 m away from mother trees (average distance between neighbouring trees was 7.4 m). Thus, insect pollinators are efficient promoters of cross-fertilization in this orchard, with the result that the effective number of males mating with each female is large.     

Somatic Cell and Sperm Cell Cytogenetics in a Patient with t(14; 21)

Approximately 15%-20% of clinically recognizable pregnancies end in spontaneous abortion. About half of the spontaneous abortions in the early stage of the pregnancy are due to chromosomal abnormalities. Using GTG chromosome banding and dual-color fluorescence in situ hybridization (FISH) techniques, we determined the cytogenetic aberration in the husband of a couple with spontaneous recurrent abortions. Karyotype analysis showed 46, XX in the wife and 45, XY, −14, −21, +t(14; 21) in the husband. We studied the mechanism of formation of the abnormal chromosome with Robertsonian translocation between chromosomes 14 and 21 by FISH and flow cytometric sorting in the sperm cells. The result showed that 71% of the gametes were balanced and the remaining 29% were not. As a result, the couple was given genetic counseling.     

Lack of effects of atomic bomb radiation on genetic instability of tandem-repetitive elements in human germ cells.

In a pilot study to detect the potential effects of atomic bomb radiation on germ-line instability, we screened 64 children from 50 exposed families and 60 from 50 control families for mutations at six minisatellite loci by using Southern blot analysis with Pc-1, lambda TM-18, ChdTC-15, p lambda 3, lambda MS-1, and CEB-1 probes. In the exposed families, one or both parents received a radiation dose > 0.01 Sv. Among the 64 children, only one child had parents who were both exposed. Thus, of a total of 128 gametes that produced the 64 children, 65 gametes were derived from exposed parents and 63 were from unexposed parents, the latter being included in a group of 183 unexposed gametes used for calculating mutation rates. The average parental gonadal dose for the 65 gametes was 1.9 Sv. We detected a total of 28 mutations at the p lambda g3, lambda MS-1, and CEB-1 loci, but no mutations at the Pc-1, lambda TM-18, and ChdTC-15 loci. We detected 6 mutations in 390 alleles of the 65 exposed gametes and 22 mutations in 1098 alleles of the 183 gametes from the unexposed parents. The mean mutation rate per locus per gamete in these six minisatellite loci was 1.5% in the exposed parents and 2.0% in the unexposed parents. We observed no significant difference in mutation rates in the children of the exposed and the unexposed parents (P = .37, Fisher's exact probability test).     

Stem cells of Hydra magnipapillata can differentiate into somatic cells and germ line cells

We investigated whether all stem cells of Hydra can differentiate both somatic cells and gametes or if a separate germ line exists in these phylogenetically old organisms. The differentiation potential of single stem cells was analyzed by applying a statistical cloning procedure. All stem cell clones were found to differentiate somatic cells. No clone was found to contain stem cells which do not differentiate. Most of the clones could be induced to form gametes. No clone was found that produced gametes only. The results indicate that stem cells are multipotent in the sense that individual stem cells can differentiate into somatic cells as well as germ line cells.     

The segregation pattern of a translocation quadrivalent

The segregation pattern of a translocation quadrivalent was studied in three hybrid families of the tetraploid (2n=28) oat, of the Avena strigosa polyploid complex. The rate of IV formation in F₁ was high and the fertility was normal. The adjacent-alternate orientation of this quadrivalent was very variable. Conspicuous variation in this frequency was found between anthers of the same floret and between florets. The alternate type was usually more common toward the end of MI. The adjacent-alternate ratio was found to be an unreliable measure for calculating the type of gametes produced by the F₁ and the F₂ plants derived from them. An attempt was made to determine the type of viable gametes produced on the F₁ and their frequency by examining the cytology of F₂ plants. The various combinations of the chromosomes of the translocation complex expected in the F₂ plants were derived and their expected frequencies calculated by taking into account chiasma formation at the chromosome ends and various restrictions of gamete viability. In none of the three hybrid families F₂ individuals were found to produce trivalents as the most complex chromosome configuration, indicating that one type of gamete derived from adjacent separation was not formed. The 11 ratio between F₂ plants having only bivalents (2II) and those with F ₁-like quadrivalent configuration R(c), expected when gametes resulting from alternate separation are fully functional, with the exclusion of other types, was not found. That ratio 2II/R(c), was 2-1/3 in the various families. The cytology of selected F₃ individuals basically followed the predictions based on chromosome association in the F₂.     

Transmission of sex cells of one species through the body of a second species in the genus Xenopus. II. Interspecific matings

Hybrids between X. mulleri and X. laevis have been prepared by two methods. By one method the hybridization has been effected either by a natural coupling following hormonal stimulation or by means of an artificial fertilization. By the other method, the primordial germ cells of mulleri have been introduced into laevis at the neurula stage and, when the host laevis had reached sexual maturity, the mulleri gametes produced have been brought into contact with laevis gametes through a natural coupling.     

Mate choice among yeast gametes can purge deleterious mutations

Meiosis in Saccharomyces yeast produces four haploid gametes that usually fuse with each other, an extreme form of self-fertilization among the products of a single meiosis known as automixis. The gametes signal to each other with sex pheromone. Better-quality gametes produce stronger signals and are preferred as mates. We suggest that the function of this signalling system is to enable mate choice among the four gametes from a single meiosis and so to promote the clearance of deleterious mutations. To support this claim, we construct a mathematical model that shows that signalling during automixis (i) improves the long-term fitness of a yeast colony and (ii) lowers its mutational load. We also show that the benefit to signalling is greater with larger numbers of segregating mutations.     

Rates of movement of transposable elements inDrosophila melanogaster

Mobilization rates of nine families of transposable elements (P, hobo, FB, gypsy, 412, copia, blood, 297, andjockey) were estimated by using 182 lines. Lines were started from a completely isogenic population ofDrosophila melanogaster, carrying the markersepia as an indicator of possible contamination, and have been accumulating spontaneous mutations independently for 80 generations of brother-sister (or two double-first-cousin) matings. Transposable element movements have been analyzed in complete genomes by the Southern technique. Mobilization was a rare event, with an average rate of 10^?5 per site per generation. The most active element wasFB. In contrast, the retroelementsgypsy andblood did not move at all. Most changes in restriction patterns were consistent with rearrangements rather than with true transposition. The euchromatic or heterochromatic location of elements was tested by comparing insertion patterns from adults and salivary glands. Certain putative rearrangements involved heterochromatic copies of the retroelements412, copia or297. Clustering of movement across families was observed, suggesting that movement of different families may be non-independent. An association between modified insertion patterns and mutant effects on quantitative traits shows that spontaneous transposition events cause continuous variation.     

Production of anisogametes and gamete motility dimorphism in Monostroma angicava

 The reproductive strategy of a marine alga with a heteromorphic biphasic life cycle was studied by analyzing various sexual reproductive characters in light of the evolution of anisogamy. Gametophytes of Monostroma angicava were dioecious and their gametes were slightly anisogamous. Volume of gametangium, density of gametangia and area of mature gametangial parts on each gametophyte did not differ from male to female. Therefore, the reproductive biomass investment for gamete production was considered to be the same for each sex. Anisogamy in this alga appeared to be derived from the difference in the number of cell divisions during gametogenesis, because the majority of male gametangia each produced 64 (2⁶) gametes and the female produced 32 (2⁵) gametes. This corresponded with measurements of cell size in male and female gametes. Further, the sex ratio was 1:1 for sexually mature plants sampled at Charatsunai. Therefore, it was suggested that in the field twice as many male gametes are released as female gametes. Liberated gametes of both sexes showed positive phototaxis. The swimming velocity of freshly liberated male gametes was a little higher than that of female gametes. Male gametes had the potential to swim for ca. 72 h and female gametes for ca. 84 h. The difference in gamete motility between the two sexes seemed to be related to cell size. Planozygotes were negatively phototactic and swam more rapidly than gametes of either sex. Received: 5 March 1997 / Revision accepted: 18 July 1997     

On sexual agglutination and mating type substances in isogamous dioecious Chlamydomonads: IV. Unilateral inactivation of the sex contact capacity in compatible and incompatible taxa by α-mannosidase and snake venom protease

Sex cell contact at fertilization is analysed in the mating type reaction of isogamous Chlamydomonas species. Contact is based upon a complementarity between special mating type substances, sex and species specific glycoprotein complexes. In three related taxa, the contact capactiy of their (+) gamete type is sensitive to snake venom protease (α-protease) and depends decisively on terminal α-glycosidically bound mannose residues. Enzymatic removal of these residues by α-mannosidase incapacitates live (+) gametes and inactivates isolated (+) mating type substance. (+) gametes inactivated by α-mannosidase or α-protease do not agglutinate with (?) gametes nor respond to isolated (?) mating type substance. Isolated (?) substance is adsorbed to and inactivated by the homologous (+) gametes. (+) gametes incapacitated by α-mannosidase or α-protease do not adsorb nor inactivate the isolated (?) substance. The agglutinability of live (?) gametes and the contact capacity of isolated (?) mating type substance is not affected by α-mannosidase or α-protease. The mannose residues react only within the species-typical complementarity. Some additional feature(s) of the (+) mating type substances must effect their species specificity and account for gametic isolation and sexual incompatibility between species.     

Mating interactions between coexisting dipoloid, triploid and tetraploid cytotypes ofHieracium Echioides (Asteraceae)

Experimental crosses between diploids, triploids and tetraploids ofHieracium echioides were made to examine mating interactions. Specifically, cytotype diversity in progeny from experimental crosses, intercytotype pollen competition as a reproductive barrier between diploids and tetraploids, and differences in seed set between intra- and intercytotype crosses were studied. Only diploids were found in progeny from 2x × 2x crosses. The other types of crosses yielded more than one cytotype in progeny, but one cytotype predominated in each cross type: diploids (92%) in 2x × 3x crosses, tetraploids (88%) in 3x × 2x crosses, triploids (96%) in 2x × 4x crosses, triploids (90%) in 4x × 2x crosses, tetraploids (60%) in 3x × 3x crosses, pentaploids (56%) in 3x × 4x crosses, triploids (80%) in 4x × 3x crosses and tetraploids (88%) in 4x × 4x crosses. No aneuploids have been detected among karyologically analyzed plants. Unreduced egg cell production was detected in triploids and tetraploids, but formation of unreduced pollen was recorded only in two cases in triploids. Triploid plants produced x, 2x and 3x gametes: in male gametes x (92%) gametes predominated whereas in female gametes 3x (88%) gametes predominated. Cytotype diversity in progeny from crosses where diploids and tetraploids were pollinated by mixture of pollen from diploid and tetraploid plants suggested intercytotype pollen competition to serve as a prezygotic reproductive barrier. No statistically significant difference in seed set obtained from intra- and intercytotype crosses between diploids and tetraploids was observed, suggesting the absence of postzygotic reproductive barriers among cytotypes.     

Analysis of gamete and zygote motility in Allomyces

To study the mechanisms of chemotaxis in eukaryotes, the motility patterns of the gametes and zygotes and the chemotactic responses of the male gametes of the lower eukaryote Allomyces macrogynus were examined. Dark-field microscopy of the male gametes showed a smooth swimming pattern interrupted by very brief ‘jerks’ of the cell body that caused a change in swimming direction. Female gametes had a slower swimming velocity than the males and underwent more jerks or turns which accounted for their sluggish motility. The zygotes swam with the fastest velocity and were observed to have a helical swimming pattern involving a continuous turning of the cell body, a behavior absent from the gametes. Introduction of female gametes that produce the chemoattractant sirenin brought about an immediate change in the behavior of the male gametes. They moved in spirals (or helices) towards the source of the chemoattractant (the female gametes), undergoing only a few jerks to reorient the male cells. When very near the female cells, or in high concentrations of added sirenin, many very short motility tracks were observed that finally resulted in contact between the two gamete types. The results indicate that the poor swimming ability of the female gametes facilitates gamete contact, resulting in as many as 30–40 male gametes clustered on a single female cell. Further, male gamete orientation to the sirenin gradient is caused by the chemoattractant suppressing the jerk motion.     

Adaptive significance of self-fertilization in a hermaphroditic perennial, Trillium camschatcense (Melanthiaceae)

The evolution of self-fertilization from primarily outcrossing ancestors is one of the most common evolutionary transitions in plants; however, the ecological mechanisms that maintain self-fertilization have remained controversial. Theoretical studies suggest that selfing is advantageous over outcrossing in terms of genetic transmission and assurance of seed production under pollen-limited circumstances. Trillium camschatcense is a herbaceous perennial distributed in Hokkaido and northern Honshu, Japan. Geographical variation in the breeding system (self-compatible, SC; or self-incompatible, SI) has been reported in populations in Hokkaido. Here, we used several SC and SI populations of T. camschatcense to investigate the adaptive significance and the evolutionary basis of self-fertilization. Pollination experiments and genetic analyses demonstrated that the potential availability of outcross pollen in SC populations was sufficient and that the number of pollen donors was equal to that of SI populations. However, despite the high availability of outcross pollen, the SC populations produced seeds predominantly by selfing and so underwent severe inbreeding depression. Although none of the suggested advantages for self-fertilization were supported by our analyses, we propose two possible scenarios for the evolution of self-fertilization in T. camschatcense.     

POLLEN DISCOUNTING AND THE SPREAD OF A SELFING VARIANT IN TRISTYLOUS EICHHORNIA PANICULATA: EVIDENCE FROM EXPERIMENTAL POPULATIONS

Floral traits that increase self-fertilization are expected to spread unless countered by the effects of inbreeding depression, pollen discounting (reduced outcross pollen success by individuals with increased rates of self-fertilization), or both. Few studies have attempted to measure pollen discounting because to do so requires estimating the male outcrossing success of plants that differ in selfing rate. In natural populations of tristylous Eichhornia paniculata, selfing variants of the mid-styled morph are usually absent from populations containing all three style morphs but often predominate in nontrimorphic populations. We used experimental garden populations of genetically marked plants to investigate whether the effects of population morph structure on relative gamete transmission by unmodified (M) and selfing variants (M‘) of the mid-styled morph could explain their observed distribution. Transmission through ovules and self and outcross pollen by plants of the M and M’ morphs were compared under trimorphic, dimorphic (S morph absent), and monomorphic (L and S morphs absent) population structures. Neither population structure nor floral morphology affected female reproductive success, but both had strong effects on the relative transmission of male gametes. The frequency of self-fertilization in the M' morph was consistently higher than that of the M morph under all morph structures, and the frequency of self-fertilization by both morphs increased as morph diversity of experimental populations declined. In trimorphic populations, total transmission by the M and M' morphs did not differ. The small, nonsignificant increase in selfing by the M' relative to the M morph was balanced by decreased outcross siring success, particularly on the S morph. In populations lacking the S morph, male gamete transmission by the M' morph was approximately 1.5 times greater than that by the M morph because of both increased selfing and increased success through outcross pollen donation. Therefore, gamete transmission strongly favored the M' morph only in the absence of the S morph, a result consistent with the distribution of the M' morph in nature. This study indicates that floral traits that alter the selfing rate can have large and context-dependent influences on outcross pollen donation.     

Test of recessive lethals in the mouse

The presence of spontaneous lethal mutations in inbred strains is discussed with special reference to their variation and influence on estimates to induced mutations.A model is presented that will facilitate classification of lethal-free and lethal heterozygotes.The model is used in classification of sons to lethal heterozygous males carrying a spontaneous mutation.The observed results are in good agreement with the model.From experience it is concluded that the most efficient way to use the facilities in lethal tests is to examine 10 or more full brothers to the P parents. By doing so pre-existing spontaneous lethals can be excludde by eliminating families in which any of the P parents were lethal heterozygous. The observed total rate of recessive lethals gives slight over-estimation of the induced rate of mutations, as spontaneous mutations in the gametes forming the F₁ cannot be excluded.