Interspecific incompatibility due to stylar barriers in tuber-bearing and closely related non-tuber-bearing Solanums

Summary The phenomenon of interspecific incompatibility between various wild tuber-bearing and closely related non-tuber-bearing Solanum species was studied. One area of investigation included an examination of possible protein interactions in the incompatibility reaction using SDS electrophoresis. Pollen tube inhibition and morphology were examined in conjunction with biochemical analysis. Two sets of crosses were examined: interspecific tuber-bearing species crosses and interspecific tuber-bearing × non-tuber-bearing species crosses. These crosses had consistent pollen tube inhibition in the upper one-third of the style. The upper third of the styles of incompatibly pollinated, compatibly pollinated, and unpollinated styles was studied under fluorescence microscopy to observe pollen tube growth and morphology. Interspecific tuber-bearing × non-tuber-bearing species crosses demonstrated consistent pollen tube inhibition just below the stigma with frequent pollen tube swelling and bursting and extensive callose deposition along the pollen tube wall. Interspecific tuber-bearing species crosses had pollen tube inhibition further down the style with pollen tube tip tapering and extensive callose deposition. Stylar proteins of the lower two-thirds of the styles were analyzed with SDS electrophoresis. No unique protein differences were found to be specifically associated with the interspecific incompatibility reaction in this portion of the style.Cooperative investigation of the U.S. Department of Agriculture, Agricultural Research Service, and the Wisconsin Experiment Station. Supported in part by the USDA, Cooperative States Research Service Competitive grant no. 83-CRCR-1-1253     

Irregular segregation at the Pr locus controlling plastid inheritance in Pelargonium: gametophytic lethal or incompatibility system?

Summary Two distinct segregation patterns are recognized after G X W plastid crosses in Pelargonium. Type I parents produce offspring in which maternal zygotes are frequent, biparental intermediate, and paternal zygotes rare (MZ>BPZ>PZ), as defined by the presence or absence of green or white plastids in the young embryos into which the zygotes develop. Type II parents produce offspring in which maternal and paternal zygotes are frequent with biparental zygotes the least frequent class (MZ>BPZPr ₁ Pr ₁. Type II plants, which do not breed true, are regarded as heterozygotes — Pr ₁ Pr ₂. The nuclear gene is symbolized as Pr as it is presumed to control alternative patterns of plastid segregation through an effect on plastid replication.Selfs and intercrosses of heterozygous plants segregate in an unexpected 1:1 ratio and not the expected 3:1 (1:2:1). The alternative homozygote — Pr ₂ Pr ₂ — could not be detected. Reciprocal crosses between heterozygotes (Pr ₁ Pr ₂) and homozygotes (Pr ₁ Pr ₁) give the expected 1:1 ratio when the Pr ₂ allele is derived from the male, whereas there is often, but not always, a highly significant deviation from 1:1 when the Pr ₂ allele is derived from the female.A simple explanation, which is not wholly satisfactory, is to assume that Pr ₂ is a gametophytic lethal on the female side. An alternative, or additional, explanation is that an incompatibility mechanism is involved in which Pr ₁ is a self-compatible allele, Pr ₂ a self-incompatible allele, and Pr ₁-Pr ₂ cross-compatible alleles. Successful fertilization is then determined by sporophytic control on the male side and gametophytic control on the female side.     

Cytoplasmic incompatibilities in the mosquito Culex pipiens: How to explain a cytotype polymorphism?*

Although cytoplasmic incompatibilities have been used as a means of eradicating the mosquito Culex pipiens, the population dynamics of these sterilities in relation to the coexistence of multiple incompatible cytotypes in a single area has not been investigated, except in the case of two unidirectionally incompatible cytotypes. An analytical model of the evolution of n cytotypes in an infinite panmictic population has been developed in order to investigate polymorphic equilibrium. A necessary criterion for the stability of such an equilibrium is established; it is shown that a stable polymorphism cannot exist between incompatible cytotypes. This result is discussed in the light of population dynamics and genetics of Culex pipiens, and of our present knowledge on incompatibilities. The consequences of a geographic structuring and of homogamy are considered. A careful reconsideration of previous experimental results disclosed probable nuclear effects and a serious experimental weakness: with the common procedure of backcrossing hybrid females to males of constant genotype it is not possible to rule out probable nuclear effects with paternal expression. It is concluded that incompatibilities in Culex pipiens may have a nuclear-cytoplasmic determinism.     

New hypothesis elucidates self-incompatibility in the olive tree regarding S-alleles dominance relationships as in the sporophytic model

Most olive varieties are not strictly self-incompatible, nevertheless, they request foreign pollen to enhance fruit yield, and consequently orchards should contain pollinisers to ensure fruit set of the main variety. The best way to choose pollinisers is to experiment numerous crosses in a diallel design. Here, the genetic mode of inheritance of SI in the olive is deciphered and it does not correspond to the GSI type, but to the SSI type. It leaves S-allele dominance relationship expression in the male (pollen and pollen tube), but not in the female (stigma and style). Thus, a pair-wise combination of varieties may be inter-compatible in one direction (male to female, or female to male) and inter-incompatible in the other direction. Dominance relationships also explain different levels of self-pollination observed in varieties. Little efficient pollinisers were found and predicted in varieties; nevertheless, some new efficient pair-wise allele combinations are predicted and could be created. This model enables one to forecast compatibility without waiting for several years of yield records and to choose pollinisers in silico.     

Promiscuous females avoid inbreeding by controlling sperm storage

Recent studies in a variety of species have shown that polyandrous females are somehow able to bias paternity against their relatives postcopulation, although how they do so remains unknown. Field crickets readily mate with their siblings, but when also mated to an unrelated male, they produce disproportionately fewer inbred offspring. We use a new competitive microsatellite polymerase chain reaction technique to determine the contribution of males to stored sperm and subsequent paternity of offspring. Paternity is almost completely predicted by how much sperm from a particular male is stored, and unrelated males contribute more sperm to storage and have a corresponding higher paternity success.     

Between- and within-host species selection on cytoplasmic incompatibility-inducing Wolbachia in haplodiploids

Abstract The most common effect of the endosymbiont Wolbachia is cytoplasmic incompatibility (CI), a form of postzygotic reproductive isolation that occurs in crosses where the male is infected by at least one Wolbachia strain that the female lacks. We revisited two puzzling features of Wolbachia biology: how Wolbachia can invade a new species and spread among populations, and how the association, once established in a host species, can evolve, with emphasis on the possible process of infection loss. These questions are particularly relevant in haplodiploid species, where males develop from unfertilized eggs, and females from fertilized eggs. When CI occurs in such species, fertilized eggs either die (female mortality type: FM), or develop into males (male development type: MD), raising one more question: how transition among CI types is possible. We reached the following conclusions: (1) the FM type is a better invader and should be retained preferentially after a new host is captured; (2) given the assumptions of the models, FM and MD types are selected on neither the bacterial side nor the host side; (3) selective pressures acting on both partners are more or less congruent in the FM type, but divergent in the MD type; (4) host and symbiont evolution can drive infection to extinction for all CI types, but the MD type is more susceptible to the phenomenon; and (5) under realistic conditions, transition from MD to FM type is possible. Finally, all these results suggest that the FM type should be more frequent than the MD type, which is consistent with the results obtained so far in haplodiploids.     

Fertility restoration and mitochondrial nucleic acids distinguish at least five subgroups among cms-S cytoplasms of maize (Zea mays L.)

Summary Differences in fertility restoration and mitochondrial nucleic acids permitted division of 25 accessions of S-type male sterile cytoplasm (cms-S) of maize into five subgroups: B/D, CA, LBN, ME, and S(USDA). S cytoplasm itself (USDA cytoplasm) was surprisingly not representative of cms-S, since only two other accessions, TC and I, matched its mitochondrial DNA pattern. CA was the predominant subgroup, containing 18 of the 25 accessions. The B/D and ME subgroups were the most fertile and LBN the most sterile. The exceptional sterility of LBN cytoplasm makes it the most promising of the 25 cms-S accessions for the production of hybrid seed. The most efficient means of quantifying the fertility of the subgroups was analysis of pollen morphology in plants having cms-S cytoplasm and simultaneously being heterozygous for nuclear restorer-of-fertility (Rf) genes. This method took advantage of the gametophytic nature of cms-S restoration. The inbred NY821LERf was found to contain at least two restorer genes for cms-S. Fertility differences were correlated with mitochondrial nucleic acid variation in the LBN, ME, and S (USDA) subgroups.Paper No. 9498 of the Journal Series of the North Carolina Agricultural Research Service, Raleigh, NC     

GENOME‐WIDE INVESTIGATION OF REPRODUCTIVE ISOLATION IN EXPERIMENTAL LINEAGES AND NATURAL SPECIES OF NEUROSPORA: IDENTIFYING CANDIDATE REGIONS BY MICROARRAY‐BASED GENOTYPING AND MAPPING

Inherent incompatibilities between genetic components from genomes of different species may cause intrinsic reproductive isolation. In evolution experiments designed to instigate speciation in laboratory populations of the filamentous fungus Neurospora, we previously discovered a pair of incompatibility loci (dfe and dma) that interact negatively to cause severe defects in sexual reproduction. Here we show that the dfe‐dma incompatibility also is a significant cause of genetic isolation between two naturally occurring species of Neurospora (N. crassa and N. intermedia). The strong incompatibility interaction has a simple genetic basis (two biallelic loci) and antagonistic epistasis occurs between heterospecific alleles only, consistent with the Dobzhansky–Muller model of genic incompatibility. We developed microarray‐based, restriction‐site associated DNA (RAD) markers that identified ～1500 polymorphisms between the genomes of the two species, and constructed the first interspecific physical map of Neurospora. With this new mapping resource, the approximate genomic locations of the incompatibility loci were determined using three different approaches: genome scanning, bulk‐segregant analyses, and introgression. These population, quantitative, and classical genetics methods concordantly identified two candidate regions, narrowing the search for each incompatibility locus to only ～2% of the nuclear genome. This study demonstrates how advances in high‐throughput, genome‐wide genotyping can be applied to mapping reproductive isolation genes and speciation research.     

Structural insights into Rhino‐Deadlock complex for germline piRNA cluster specification

PIWI‐interacting RNAs (piRNAs) silence transposons in germ cells to maintain genome stability and animal fertility. Rhino, a rapidly evolving heterochromatin protein 1 (HP1) family protein, binds Deadlock in a species‐specific manner and so defines the piRNA‐producing loci in the Drosophila genome. Here, we determine the crystal structures of Rhino‐Deadlock complex in Drosophila melanogaster and simulans. In both species, one Rhino binds the N‐terminal helix–hairpin–helix motif of one Deadlock protein through a novel interface formed by the beta‐sheet in the Rhino chromoshadow domain. Disrupting the interface leads to infertility and transposon hyperactivation in flies. Our structural and functional experiments indicate that electrostatic repulsion at the interaction interface causes cross‐species incompatibility between the sibling species. By determining the molecular architecture of this piRNA‐producing machinery, we discover a novel HP1‐partner interacting mode that is crucial to piRNA biogenesis and transposon silencing. We thus explain the cross‐species incompatibility of two sibling species at the molecular level.