Variable responses of insects to hybrid versus parental sagebrush in common gardens

Both ecological and genetic mechanisms have been proposed to explain patterns of herbivore attack on interspecific plant hybrids, but distinguishing among them can be difficult in natural hybrid zones. We performed a common-garden experiment to evaluate four genetic hypotheses: dominance, additivity, elevated hybrid susceptibility, and elevated hybrid resistance. Censuses and cage experiments were used to compare insect responses to basin big sagebrush (Artemisia tridentata spp. tridentata), mountain big sagebrush (A. t. vaseyana), and their F₂ progeny. After two growing seasons, hybrid shrubs resembled mountain big sagenbrush in size, but were more similar to basin big sagebrush in flower production. Censuses of naturally colonizing insects (the gall midge Rhopalomyia obovata, the bagworm moth Apterona helix, and the aphid Obtusicauda coweni) tended to support the dominance hypothesis: if the insect clearly discriminated between the two parents, its frequency on hybrids closely resembled that on one of the parents. Moreover, colonization of hybrids in all three cases suggested a dominance deviation toward the susceptible parent rather than toward the resistant parent. In contrast to the censuses, cage experiments involving two insects supported the hybrid-susceptibility hypothesis; both survival and growth of the grasshopper Melanoplus sanguinipes and growth of the leaf beetle Trirhabda pilosa were higher on hybrid shrubs than on either parent. Because many secondary compounds have been determined to occur at intermediate concentrations in F₂ shrubs, dominance for susceptibility may indicate that insects respond to plant traits (e.g., oviposition stimulants and deterrents) in a threshold manner. Mechanisms underlying increased hybrid susceptibility are less clear, but our experimental design makes environmental explanations (e.g., the plant-stress hypothesis) unlikely. Although we eliminated several confounding factors, our results agree with the conclusion from natural hybrid zones that insect responses to hybrid plants are likely to be idiosyncratic; even congeneric species did not respond similarly to hybrid and parental plants.     

Nature and type of gene action governing resistance to Helminthosporium turcicum leaf blight in maize (Zea mays L.)

Six generations, consisting of three resistant parents, three susceptible parents, their 15 possible F₁ crosses, 15 F₂'s, 15 BC₁'s (F₁ x resistant female parent) and 15 BC₂'s (F₁ x susceptible male parent) were analysed following Hayman (Heredity 12: 371–390, 1958) to evaluate the nature and type of gene action governing resistance to H. turcicum. The results showed that all types of gene effects, viz., additive, dominance and epistasis (i.e., additive x additive, additive x dominance and dominance x dominance) were operating in one cross or the other in controlling resistance. However, it was additive gene action and dominance x dominance type of epistasis with duplicate nature that were important in controlling resistance in most crosses. Depending upon the final objectives, one of the breeding methods, viz., recurrent selection, heterosis breeding, back-cross method or full-sib selection (bi-parental mating) may be followed.     

Quantitative genetics of growth and development in Populus. I. A three-generation comparison of tree architecture during the first 2 years of growth

One approach to gain an insight into the genetics of tree architecture is to make use of morphologically divergent parents and study their segregating progeny in the F₂ and backcross (B₁) generations. This approach was chosen in the present study in which material of a three-generation pedigree growing side by side in a replicated plantation, was analyzed. The pedigree included Populus trichocarpa (T) and P. deltoides (D) parents, their F₁ and F₂ hybrids and their B₁ hybrids to the D parent. The trees were grown in the environment of the T parent and measured for the first 2 years of growth. Nine quantitative traits were studied at the stem, branch and leaf levels of tree architecture, in which the original parents differed. Strong F₁ hybrid vigor relative to the better parent (T) was expressed in growth and its components. Most quantitative traits in the F₂ and B₁ hybrids were intermediate between the T and D parents but displayed a wide range of variation due to segregation. The results from the analysis of variance indicated that all morphometric traits were significantly different among F₂ and B₁ clones, but the B₁ hybrids were more sensitive to replicates than the F₂. Broad-sense heritabilities (H ²) based on clonal means ranged from moderately high to high (0.50–0.90) for the traits studied, with H ² values varying over age. The H ² estimates reflected greater environmental noise in the B₁ than in the F₂, presumably due to the greater proportion of maladaptive D alleles in those hybrids. In both families, sylleptic branch number and length, and leaf size on the terminal, showed strong genetic correlations with stem growth. The large divergence between the two original parents in the traits studied, combined with the high chromosome number in Populus (2n=38), makes this pedigree well suited for the estimation of the number of quantitative trait loci (QTLs) underlying quantitative variation by Wright's biometric method (1968). Variation in several traits was found to be under the control of surprisingly few major QTLs: 3–4 in 2nd-year height and diameter growth, a single QTL in stem diameter/height ratio.     

超甜玉米种子萌发物质利用性状杂种优势及亲子回归关系分析

为阐明超甜玉米(Zea mays L.var.saccharata Bailey)亲本对F_1种子物质利用性状遗传效应,研究了19份自交系和其测配的10个杂交组合的杂种优势及亲子回归关系。结果表明,超甜玉米自交系及F_1种子的淀粉含量、蛋白质含量、脂肪含量、百粒重、种子物质动用量和种子物质利用率差异较大,10个杂交组合中亲本和F_1种子的淀粉含量、脂肪含量、百粒重均存在显著差异。F_1种子淀粉含量和百粒重均表现出正向超亲优势,即近高亲本遗传;而F_1种子的蛋白质含量、脂肪含量、种子物质动用量和种子物质利用率为近低亲本遗传。聚类分析和杂种优势分析表明,性状差异较大的FH14、Q26、GT22、GT2等亲本测配的杂交组合在种子百粒重或种子物质利用率等性状上具有较强的超亲优势。回归分析表明,母本对F_1种子的淀粉含量、百粒重有负效应,对种子物质动用量和种子物质利用率有正效应;父本对种子淀粉含量有负效应,对种子物质利用率有正效应。     

Inheritance of the photoperiodically induced cold acclimation response in Cornus sericea L., red-osier dogwood

Summary Cold acclimation responses of latitudinal ecotypes of Cornus sericea L. (C. stolonifera Michx.) and F₁, F₂ and BC₁ hybrid progenies were measured under natural photoperiod conditions in St. Paul, MN and artificially shortened photoperiods in the glass-house. The 65 °N and 62 °N ecotypes (Alaska and Northwest Territories, respectively) were characterized by a short night length for hardiness induction, the 42 °N ecotype (Utah A and B) by a long night length for hardiness induction, while the F₁ was intermediate to the parents. Results from reciprocal crosses indicated there was no significant unilateral maternal influence on cold acclimation. Acclimation responses of the F₂ were highly variable but generally ranged between the parental extremes. However, three individuals from the 42 ° × 62 °N crosses exhibited greater cold resistance than the northern parent on two successive freezing test dates. F₂ plants were also found with less freezing resistance than the southern parent. Backcrosses to the southern parent produced progeny with acclimation patterns resembling that of the southern parent and were significantly less hardy than the F₂ in early freezing tests.Scientific Journal Series Paper No. 12,075 of the Minnesota Agricultural Experiment Station     

Inheritance of grain filling duration in spring wheat (<Emphasis Type="Italic">Triticum aestivum</Emphasis> L. em thell)

To understand the genetic control of grain filling duration (GFD), i.e., the number of days from anthesis to physiological maturity, we studied the F₁, F₂, BC1 and BC₂ generations of six spring wheat crosses from nine varieties/genotypes. Generation mean analysis for gene effects indicated that one or more types of epistasis were significant in all crosses. In each pairing, the F₁ and F₂ means were either intermediate or closer to the mean of the parent having the longer GFD. Our narrow-sense heritability estimate was reasonably high, at 47.67 (based on diallel analysis). This demonstrated that progress could be made from the selection in these crosses for either long or short GFD. The two early varieties that had identical maturity durations differed in their GFD values, indicating that maturity dates are not good criteria when choosing parents for modifying GFD. To utilize favorable additive × additive effects during this selection, we suggest that a single seed descent (SSD) or bulk popula-tion approach be adopted. In comparison, dominance effects would prove quite useful in hybrid wheat breeding programs.     

ECOLOGICAL SPECIATION IN STICKLEBACKS: ENVIRONMENT-DEPENDENT HYBRID FITNESS

“Ecological” speciation occurs when reproductive isolation evolves as a consequence of divergent selection between populations exploiting different resources or environments. We tested this hypothesis of speciation in a young stickleback species pair by measuring the direct contribution of ecological selection pressures to hybrid fitness. The two species (limnetic and benthic) are strongly differentiated morphologically and ecologically, whereas hybrids are intermediate. Fitness of hybrids is high in the laboratory, especially F₁ and F₂ hybrids (backcrosses may show some breakdown). We transplanted F₁ hybrids to enclosures in the two main habitats in the wild to test whether the distribution of resources available in the environment generates a hybrid disadvantage not detectable in the laboratory. Hybrids grew more slowly than limnetics in the open water habitat and more slowly than benthics in the littoral zone. Growth of F₁ hybrids was inferior to the average of the parent species across both habitats, albeit not significantly. The contrast between laboratory and field results supports the hypothesis that mechanisms of F₁ hybrid fitness in the wild are primarily ecological and do not result from intrinsic genetic incompatibilities. Direct selection on hybrids contributes to the maintenance of sympatric stickleback species and may have played an important role in their origin.     

Genetic studies of yield contributing traits in Amaranthus

Summary Genetical studies on grain yield and its contributing traits were made in a six parent complete diallel in the F₁ and F₂ generations of one of the most widely grown grain species of grain amaranths (Amaranthus hypochondriacus L.). Graphical analysis indicated that epistasis exists for 1,000-grain weight in the F₁. Grain weight/panicle, yield/plant and harvest index indicated absence of non-allelic gene interaction. The harvest index in the F₁ and F₂ and grain weight/ panicle, 1,000-grain weight, yield/plant in the F₂ appeared to be controlled by overdominance effects. Higher grain yield appeared to be associated with dominant genes. Both additive and non-additive gene effects were responsible for the genetic variation in the diallel population. However, dominance variance was more important than additive variance in grain yield/ plant and harvest index in the F₁ and F₂. For 1,000-grain weight additive genetic variance was more important in the F₁ and non-additive in F₂. There was overdominance of a consistent nature in the two analyses for harvest index in the F₁ and F₂, grain weight/ panicle, 1,000-grain weight and yield/plant in the F₂ and partial dominance for 1,000-grain weight in the F₁.     

Inheritance of resistance to leaf and glume blotch caused by Septoria nodorum Berk. in winter wheat

Sixteen crosses between eight winter wheat cultivars were screened for resistance to Septoria nodorum leaf and glume blotch in the F₁ and F₄ generations using artificial inoculation in the field. The F₁ of most crosses showed dominance for susceptibility on both ear and leaf. The effects of general combining ability were of similar magnitude as the effects for specific combining ability. On the basis of the phenotypic difference of the parents, no prediction was possible about the amount and the direction of genetic variance in the segregating populations. The variation observed in this study both within and among the segregating populations suggests a quantitative inheritance pattern influencing the expression of the two traits. The components of variance between F₂ families within a population were as high as (for S. nodorum blotch on the ear) or higher (for S. nodorum blotch on the leaf) than those between populations. Therefore, strong selection within a few populations may be as effective to obtain new resistant genotypes as selection in a large number of populations. In almost all crosses, progenies were found that were more resistant than the better parent. Thus transgression breeding may be a tool to breed for higher levels of resistance to S. nodorum blotch. Highly resistant genotypes were found even in combination with two susceptible parents. The genetic source for Septoria resistance is probably broader than is generally assumed and could be used to improve S. nodorum resistance by combination breeding followed by strong selection in large populations. Received: 18 January / Accepted: 30 April 1999     

Inheritance of components of partial resistance of barley to Rhynchosporium secalis with particular reference to race specificity

Six spring barley cultivars with no known genes for resistance to specific virulences but varying in partial resistance to Rhynchosporium secalis, were crossed in all combinations (6 × 6 diallel including reciprocals). In addition to seeds from naturally selfed plants, seeds of all parent cultivars were also produced by artificial selfing (emasculation followed by pollination using pollen from the same cultivar). This ensured comparability between seeds of parents and F₁. Both sets of parents, F₁ and F₂ families were grown in the field as single spaced plants and inoculated at Zadoks growth stage 49 with spore suspensions (2 × 10⁶ spores ml^-1) of three races (pathotypes) of R. secalis (Zadoks, Chang & Konzak, 1974). Components of partial resistance, incubation period (ICP), infection frequency (IF) and spore production per lesion (SP/L) were assessed on each plant. There were highly significant differences for all three components of partial resistance in both sets of parent cultivars but rank order in both sets was similar as evidenced by correlation coefficients, r= 0.96 for ICP and IF and r= 0.87 for SP/L. All three components of partial resistance were strongly correlated with NIAB (National Institute of Agricultural Botany, Cambridge, UK) resistance ratings. Means of F₁ and F₂ families were correlated with mid-parent values for ICP and IF but not SP/L. No difference in aggressiveness was found between races but for each component of partial resistance there was a significant interaction between race and parent cultivar (artificial selfs) and, for IF and ICP, a significant interaction between race and F₁ family. There was no evidence of interaction between parent (natural selfs) and race nor between race and F₂ family. Examination of genetic control of resistance showed evidence of strong additive effects (combining ability) in both F₁ and F₂ for ICP and IF but not for SP/L. There was no evidence for maternal or reciprocal differences, but there was evidence for dominance effects although their nature differed between components of partial resistance and between F₁ and F₂ generations. In the F₁, but not the F₂ generation, several elements of dominance (direction, distribution of dominant genes between parent cultivars, specific combining ability) showed for ICP or IF (but not SP/L) significant interaction with race.     

Chromosomal inheritance of parental rDNAs distribution pattern detected by FISH in diploid F1 hybrid progeny of Cobitis (Teleostei,Cobitidae) species has non-Mendelian character

This study was conducted to describe the major and the minor rDNA chromosome distribution in the spined loach Cobitis taenia (2n = 48) and the Danubian loach Cobitis elongatoides (2n = 50), and their laboratory-produced diploid reciprocal F₁ hybrid progeny. It was tested by fluorescence in situ hybridisation (FISH) whether the number of 28s and 5s rDNA sites in the karyotypes of diploid hybrids corresponds to the expectations resulting from Mendelian ratio and if nucleolar organiser regions (NOR)were inherited from both parents or nucleolar dominance can be observed in the induced F₁ hybrid progeny. Ten (females) or twelve (males) 28s rDNA loci were located in nine uniarm chromosomes of C. taenia. Two of such loci terminally bounded on one acrocentric chromosome were unique and indicated as specific for this species. Large 5s rDNA clusters were located on two acrocentric chromosomes. In C. elongatoides of both sexes, six NOR sites in terminal regions on six meta-submetacentric chromosomes and two 5s rDNA sites on large submetacentrics were detected. The F₁ hybrid progeny (2n = 49) was characterised by the intermediate karyotype with the sites of ribosome synthesis on chromosomes inherited from both parents without showing nucleolar dominance. 5s rDNA sites were detected on large submetacentric and two acrocentric chromosomes. The observed number of both 28s and 5s rDNAs signals in F₁ diploid Cobitis hybrids was disproportionally inherited from the two parental species, showing inconsistency with the Mendelian ratios. The presented rDNA patterns indicate some marker chromosomes that allow the species of the parental male and female to be recognised in hybrid progeny. The 5s rDNA was found to be a particularly effective diagnostic marker of C. elongatoides to partially discern genomic composition of diploid Cobitis hybrids and presumably allopolyploids resulting from their backcrossing with one of the parental species. Thus, the current study provides insight into the extent of rDNA heredity in Cobitis chromosomes and their cytotaxonomic character.     

Cytogenetics of the F1 hybrid between cassava and ceara rubber,and its backcross

Cytogenetical studies of the F₁ hybrid between the commercially cultivated tuber crop, cassava (Manihot esculenta Crantz.) and the closely related wild speciesManihot glaziovii Muell. (Ceara rubber) used as donor specles for Cassava mosaic discase and drought-resistant genes and back crosses (to cassava parent) were made. The contrasting parental characters showed partial to total dominance in the F₁ hybrid, while the back cross plants were similar to cassava in most of their characters. Eleven of the twelve backeross plants exhibited resistance to Cassava mosaic under field conditions. Karyological similarities and differences as resolved on the basis of a comparative study of the karyotypes of the cassava parent and coara rubber were corroborated by the study of chromosomal pairing in the F₁ at pachytene. Major chromosomal differentiation in the two species involved three chromosomes of their haploid complement which were represented by three heteromorphic bivalent associations in F₁ each consisting of a probably basic chromosomal type and a derived type. Pachytene analyses of three back cross plants provided direct proof for random transmission of marker chromosomes of both the parents through male gametes of the F₁ hybrid. An increase in the chiasma frequency in the back cross plants over the F₁ hybrid at metaphase I stage was also observed. Pollen fertility of the backeross plants showed considerable variation.     

Relationships of differential gene expression in leaves with heterosis and heterozygosity in a rice diallel cross

Using differential display analysis, we assessed the patterns of differential gene expression in hybrids relative to their parents in a diallel cross involving 8 elite rice lines. The analysis revealed several patterns of differential expression including: (1) bands present in one parent and F₁ but absent in the other parent, (2) bands observed in both parents but not in the F₁, (3) bands occurring in only one parent but not in the F₁ or the other parent, and, (4) bands detected only in the F₁ but in neither of the parents. Relationships between differential gene expression and heterosis and marker heterozygosity were evaluated using data for RFLPs, SSRs and a number of agronomic characters. The analysis showed that there was very little correlation between patterns of differential expression and the F₁ means for all six agronomic traits. Differentially expressed fragments that occurred only in one parent but not in the other parent or in F₁ in each of the respective crosses were positively correlated with heterosis and heterozygosity. And conversely, fragments that were detected in F₁s but in neither of the respective parents were negatively correlated with heterosis and heterozygosity. The remaining patterns of differential expression were not correlated with heterosis or heterozygosity. The relationships between the patterns of differential expression and heterosis observed in this study were not consistent with expectations based on dominance or overdominance hypotheses.     

The hybrid protein interactome contributes to rice heterosis as epistatic effects

Heterosis is the phenomenon in which hybrid progeny exhibits superior traits in comparison with those of their parents. Genomic variations between the two parental genomes may generate epistasis interactions, which is one of the genetic hypotheses explaining heterosis. We postulate that protein?protein interactions specific to F₁ hybrids (F₁‐specific PPIs) may occur when two parental genomes combine, as the proteome of each parent may supply novel interacting partners. To test our assumption, an inter‐subspecies hybrid interactome was simulated by in silico PPI prediction between rice japonica (cultivar Nipponbare) and indica (cultivar 9311). Four‐thousand, six‐hundred and twelve F₁‐specific PPIs accounting for 20.5% of total PPIs in the hybrid interactome were found. Genes participating in F₁‐specific PPIs tend to encode metabolic enzymes and are generally localized in genomic regions harboring metabolic gene clusters. To test the genetic effect of F₁‐specific PPIs in heterosis, genomic selection analysis was performed for trait prediction with additive, dominant and epistatic effects separately considered in the model. We found that the removal of single nucleotide polymorphisms associated with F₁‐specific PPIs reduced prediction accuracy when epistatic effects were considered in the model, but no significant changes were observed when additive or dominant effects were considered. In summary, genomic divergence widely dispersed between japonica and indica rice may generate F₁‐specific PPIs, part of which may accumulatively contribute to heterosis according to our computational analysis. These candidate F₁‐specific PPIs, especially for those involved in metabolic biosynthesis pathways, are worthy of experimental validation when large‐scale protein interactome datasets are generated in hybrid rice in the future.     

The cytogenetics of a Triticum turgidum x Psathyrostachys juncea hybrid and its backcross derivatives

Psathyrostachys juncea (2n = 2x = 14, NN), a source of barley yellow dwarf (BYDV) virus resistance with tolerance to drought and salinity, has been successfully hybridized in its autotetraploid form (2n = 4x = 28, NNNN) as the pollen parent to durum wheat (Triticum turgidum L.). The 2n = 4x = 28 (ABNN) F₁ hybrid has a mean meiotic metaphase-I configuration of 20.29 univalents + 0.29 ring bivalents + 3.36 rod bivalents + 0.14 trivalents. Spike length, internode length, glume awn length and lemma awn length, as well as the general spike morphology of the F₁ hybrid, are intermediate with those of the two parents. Pollinating the ABNN F₁ hybrid has given backcross (BC)-I derivatives of an amphiploid (AABBNN) that expresses limited self-fertility. BC-2 derivatives have been obtained from these plants. Direct transfers of useful genes from Ps. juncea to wheat would require substantial genetic manipulation strategies. Both conventional and novel methodologies, which may complement each other, and so facilitate reaching an agricultural objective end point, are addressed.     

Studies on trisomics in a Jute hybrid

Fifteen red pigmented trisomics were isolated in the F₂ generation from the cross Corchorus olitorius L. x C. capsularis L. In the F₃ generation a few green trisomics were obtained; more of these were isolated from the backcross generation. A detailed morphological and cytological analysis of the trisomic hybrid populations derived from the F₃ and F₄ generations is presented. The trisomics were scored for several morphologically differentiating characters and most of them were intermediate between the parental species, a few resembling the olitorius parent more. Cytological studies showed the formation of abnormal sporads in trisomics with different frequencies indicating a misdivision at meiosis. This imbalance at meiosis results in a higher percentage of pollen sterility in the trisomics as compared with the parents. Analysis of M₁ of meiosis showed that there were: differences in the frequencies of the various chromosome configurations between the two categories of trisomics; significantly higher trivalent frequencies per PMC in the green trisomics; in contrast significantly hihger univalent frequencies per PMC in the red trisomics. No significant difference in chiasma formation was observed between red and green trisomics, nor between trisomics and their parental species. It appears that segmental homology in the parental chromosomes has probably resulted in varying degrees of preferential pairing in the trisomic hybrid.     

Comparison of phenotypic and molecular distances to predict heterosis and F1 performance in Ethiopian mustard (Brassica carinata A. Braun)

Predicting heterosis and F₁ performance from the parental generation could largely enhance the efficiency of breeding hybrid or synthetic cultivars. This study was undertaken to determine the relationship between parental distances estimated from phenotypic traits or molecular markers with heterosis, F₁ performance and general combining ability (GCA) in Ethiopian mustard (Brassica carinata). Nine inbred lines representing seven different geographic regions of Ethiopia were crossed in half-diallel. The nine parents along with their 36 F₁s were evaluated in a replicated field trail at three locations in Ethiopia. Distances among the parents were calculated from 14 phenotypic traits (Euclidean distance, ED) and 182 random amplified polymorphic DNA (RAPD) markers (Jaccard’s distances, JD), and correlated with heterosis, F₁ performance and GCA sum of parents (GCA_sum). The correlation between phenotypic and molecular distances was low (r=0.34, P≤0.05). Parents with low molecular distance also had low phenotypic distance, but parents with high molecular distance had either high, intermediate or low phenotypic distance. Phenotypic distance was highly significantly correlated with mid-parent heterosis (r=0.53), F₁ performance (r=0.61) and GCA (r=0.79) for seed yield. Phenotypic distance was also positively correlated with (1) heterosis, F₁ performance and GCA for plant height and seeds plant⁻¹, (2) heterosis for number of pods plant⁻¹, and (3) F₁ performance for 1,000 seed weight. Molecular distance was correlated with GCA_sum (r=0.36, P≤0.05) but not significantly with heterosis and F₁ performance for seed yield. For each parent a mean distance was calculated by averaging the distances to the eight other parents. Likewise, mean heterosis was estimated by averaging the heterosis obtained when each parent is crossed with the other eight. For seed yield, both mean ED and JD were significantly correlated with GCA (r=0.90, P≤0.01 for ED and r=0.68, P≤0.05 for JD) and mean heterosis (r=0.79, P≤0.05 for ED and r=0.77, P≤0.05 for JD). In conclusion, parental distances estimated from phenotypic traits better predicted heterosis, F₁ performance and GCA than distances estimated from RAPD markers. Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

NUMERICAL TAXONOMY APPLIED TO CUCURBITA RELATIONSHIPS

In a numerical study of taxonomic relationships in Cucurbita, the inclusion of F₁ interspecific hybrids did not materially alter the species grouping in the phenogram. The F₁ interspecific hybrids tend to cluster with one of the parent species or species groups unless the parents are widely divergent. Also, the F₁ interspecific hybrids between wild and domesticated species cluster with the wild parent Both parent species or species groups of an F₁ interspecific hybrid can usually be determined from Q-correlation values. It is essential that compatibility relations be determined between species; i.e., species groups must be established before Q-correlation values can be successfully used in determining the parentage of hybrids. Undescribed collections can usually be correctly placed in the proper species group.     

Proteomic analysis of F1 hybrids and intermediate variants in a Littorina saxatilis hybrid zone

Proteomic analysis was carried out on the Crab (upper-shore) and Wave (lower-shore) ecotypes of Littorina saxatilis from a hybrid zone at Silleiro Cape, Spain. Proteome profiles of individual snails were obtained. Protein expression in F₁ hybrid snails bred in the laboratory and snails with intermediate shell phenotypes collected from the mid-shore were compared with Crab and Wave ecotypes using analytical approaches used to study dominance. Multivariate analysis over many protein spots showed that the F₁ snails are distinct from both ecotypes but closer to the Wave ecotype. The intermediate snails are highly variable, some closer to the Crab and others to the Wave ecotype. Considered on a protein by protein basis, some proteins are significantly closer in expression to the Crab and others to the Wave ecotype for both F₁ and intermediate snails. Furthermore, a significant majority of proteins were closer in expression to the Wave ecotype for the F₁, consistent with the multivariate analysis. No such significant majority toward either the Crab or Wave ecotype was observed for the intermediate snails. The closer similarity of F₁ and Wave ecotype expression patterns could be the result of similar selective pressures in the similar mid-shore and low-shore environments. For a significantly larger number of proteins, intermediate snails were closer in expression to the ecotype having the lower expression, for both Crab and Wave ecotypes. This is somewhat unexpected as lower expression might be expected to be an indication of impairment of function and lower fitness. Proteomic analysis could be important for the identification of candidate proteins useful for gaining improved understanding of adaptation and barriers to gene flow in hybrid zones.