Biochemical genetics of hybrid sunfish: Differential survival of heterozygotes

Allelic segregation in reciprocal backcrosses involving the largemouth bass (Micropterus salmoides) and the F₁ hybrid (largemouth bass × smallmouth bass, M. dolomieui) was investigated to determine the extent of euheterosis and luxuriance. The frequencies of allelic isozymes encoded in the lactate dehydrogenase E, malate dehydrogenase B, and isocitrate dehydrogenase loci were determined for reciprocal backcross progeny subjected to different selection pressures. The progeny of the backcross (male F₁ × female largemouth bass) underwent a rapid loss of heterozygous individuals in a natural pond environment. When the offspring of this same mating were placed in artificial pools, where cannibalism is the main source of mortality, heterozygosity was advantageous. There was a marked correlation of increased heterozygosity at these enzyme loci with an increased growth rate. None of the above responses to selection was observed when the F₁ hybrid served as the maternal parent in the reciprocal backcross. A maternal factor in the egg cytoplasm may influence the expression of heterosis.     

Inheritance patterns of enzymes and serum proteins of mallard-black duck hybrids

From 1974 to 1976, a breeding program was used to produce hybrids of black ducks and mallards for the evaluation of inheritance patterns of serum proteins and serum, liver and muscle enzymes. In addition to the crosses designed to produce hybrids, a series of matings in 1975 and 1976 were designed to evaluate inheritance patterns of a hybrid with either a black duck or mallard. At the F₁ level, hybrids were easily distinguished using serum proteins. However, once a hybrid was crossed back to either a mallard or black duck, only 12–23% of the progeny were distinguishable from black ducks or mallards using serum proteins and 23–39% using esterases. Muscle, serum and liver enzymes were similar between the two species.     

Isozyme gene markers in the dioecious species Asparagus officinalis L.

Summary Extracts from phylloclads of Asparagus officinails were electrophoretically analyzed for isozyme polymorphism. Fourteen enzyme systems were examined using four buffer systems: seven enzymes (acid phosphatase, catalase, glutamate-oxaloacetate transaminase, isocitrate dehydrogenase, malate dehydrogenase, peroxidase, and 6-phosphogluconate dehydrogenase) exhibited clear and consistent banding patterns. Isozyme polymorphism was studied in seven pairs of male and female doubled haploids and in their male F₁s. Segregation of polymorphic loci was examined in the backcross progenies and was found to be consistent with a simple Mendelian inheritance in all cases, except for three anodical peroxidases, where two factors have been hypothesized. No linkage could be found between isozyme markers that were segregating in the same cross, but association was demonstrated between one malate dehydrogenase locus and the sex determining genes. The availability of isozyme markers may be useful in breeding and, in particular, the localization of one malate dehydrogenase locus on the sex chromosomes may be helpful in mapping the sex genes.     

Inheritance of Cry1Ac resistance and associated biological traits in the cotton bollworm, Helicoverpa armigera (Lepidoptera: Noctuidae)

The analysis of reciprocal genetic crosses between resistant Helicoverpa armigera strain (BH-R) (227.9-fold) with susceptible Vadodara (VA-S) strain showed dominance (h) of 0.65-0.89 and degree of dominance (D) of 0.299-0.782 suggesting Cry1Ac resistance as a semi-dominant trait. The D and h values of F₁ hybrids of female resistant parent were higher than female susceptible parent, showing maternally enhanced dominance of Cry1Ac resistance. The progeny of F₂ crosses, backcrosses of F₁ hybrid with resistant BH-R parent did not differ significantly in respect of mortality response with resistant parent except for backcross with female BH-R and male of F₁ (BH-R × VA-S) cross, suggesting dominant inheritance of Cry1Ac resistance. Evaluation of some biological attributes showed that larval and pupal periods of progenies of reciprocal F₁ crosses, backcrosses and F₂ crosses were either at par with resistant parent or lower than susceptible parent on treated diet (0.01 μg/g). The susceptible strain performed better in terms of pupation and adult formation than the resistant strain on untreated diet. In many backcrosses and F₂ crosses, Cry1Ac resistance favored emergence of more females than males on untreated diet. The normal larval period and the body weight (normal larval growth) were the dominant traits associated with susceptible strain as contrast to longer larval period and the lower body weight (slow growth) associated with resistance trait. Further, inheritance of larval period in F₂ and backcross progeny suggested existence of a major resistant gene or a set of tightly linked loci associated with Cry1Ac sensitivity.     

Extensive heterozygosity at three enzyme loci in hybrid sunfish populations

Hybrid populations of sunfishes were produced in two different ponds, and the frequencies of allelic isozyme phenotypes were determined for three enzyme systems—malate dehydrogenase (NAD), esterases, and tetrazolium oxidase—in order to estimate the extent of heterozygosity at four different genetic loci. Interspecific F₁ hybrid fry (red-ear male × bluegill female) were produced in vitro. These fry were stocked in ponds at the free-swimming stage. When 1 year old, the F₁ hybrids produced a large F₂ hybrid population. Successful hybrid reproduction occurred each year thereafter. In one pond, a 1-year-old F₂ population exhibited all three isozyme phenotypes (red-ear, F₁, bluegill) at most loci in the approximate ratio of the 1:2:1 expected. In a second pond, 5-year-old individuals of the F₂ generation were morphologically like the F₁ and were all heterozygous for the enzyme loci studied. This unusual degree of heterozygosity in the older F₂ population appeared to be the result of differential survival of mature heterozygous individuals and not the result of early embryonic lethality. The increased heterozygosity at these unlinked loci was assumed to reflect the condition at other genetic loci in the F₂ hybrids. Several possible mechanisms are advanced to explain this apparent heterosis.This research was supported by NSF grant GB-16425 (G.S.W.) and by funds from the Illinois Natural History Survey (W.F.C.).     

Linkage relationships of six enzyme Loci in interspecific sunfish hybrids (genus lepomis)

Backcross hybrids produced from the bluegill, the red-ear sunfish, and their F₁ interspecific hybrid have been analyzed for the inheritance of six enzyme phenotypes. Malate dehydrogenase A and B, tetrazolium oxidase, 6-phosphogluconate dehydrogenase, skeletal muscle esterase, and liver α-glycerophosphate dehydrogenase are all inherited in a mendelian manner as codominant alleles at nuclear loci. 6-phosphogluconate dehydrogenase and α-glycerophosphate dehydrogenase are encoded by linked loci, undergoing recombination at a frequency of 15%-22%. No other case of linkage was observed. The absence of linkage between the homologous malate dehydrogenase loci is of particular interest. These interspecific hybrids appear to be very useful for studies of biochemical genetics.     

The inheritance of a defective lipopolysaccharide response locus in C3H/HeJ mice

F₁ hybrid mice from crosses between the lipopolysaccharide (LPS) nonresponder strain C3H/HeJ and a variety of LPS responder strains show quantitative or codominant inheritance of mitogenic responsiveness to LPS. Backcross segregation ratios indicate that a defect at a single locus is responsible for the lack of responsiveness in C3H/HeJ mice. Autoradiographic studies show that the number of LPS-responsive cells in F₁ cultures is approximately half the number of responsive cells in parent cultures. Kinetic patterns of mitogenic responsiveness to LPS differ in F₁ hybrid and parent cultures. The kinetic patterns of responsiveness vary with the cell concentrations of the culture and appear to correlate with the number of LPS-responsive cells in F₁ and parent cultures.     

Etude comparative d'activites enzymatiques microsomiques dans les hepatocytes de singe adulte et foetal: Comparative study of microsomal enzymic activities in adult and foetal monkey hepatocytes

Differential centrifugation was applied to adult and foetal liver of monkey. Obtained fractions were: F₁ (800 × g); F₂ (12 500 × g); F₃ (200 000 × g); and cell sap. Analysis of chemical compounds of these fractions shows that: (1) adult and foetal nucleic acids levels are similar; (2) there are more proteins in adult than in foetal hepatocytes; (3) most of the glycogen is located in F₃; the foetal level is twenty times higher than the adult level.Plasma membrane enzymes (5′-nucleotidase, adenylate cyclase) show a nucleomicrosomic distribution. The distribution of alkaline phosphatase is not significant.Mitochondrial enzymes (monoamine oxydase, succinate cytochrome c reductase, cytochrome oxydase) are enriched in F₂ without any sedimentation in F₃ There is more malate dehydrogenase liberated in cell sap during foetal liver fractionation. This indicates the foetal mitochondria are more sensitive to the homogenisation method.Lysosomal enzymes (acid phosphatase, N-acetylglucosaminidase) are enriched in F₂. The same observation for N-acetylglucosaminidase as for malate dehydrogenase leads to the same conclusion for foetal lysosomes.Endoplasmic reticulum and Golgi enzymes (glucose-6-phosphatase and related phosphotransferase activity, NADPH-cytochrome c reductase and sialyltransferase) are much enriched in F₃. Thus this fraction F₃ is pure enough to allow the observation of the modification produced on endoplasmic reticulum and Golgi apparatus during foetal and neonatal development.     

Inheritance and linkage analysis of five enzyme loci in interspecific hybrids of toadlets,genus Bombina

Progeny produced from Bombina bombina, B. variegata, and field-collected interspecific hybrids have been analyzed for the inheritance of five enzyme loci, which are fixed for alternate alleles in the parental species. Lactate dehydrogenase (Ldh-1), NAD-dependent malate dehydrogenase (Mdh-1), creatine kinase (Ck), adenylate kinase (Ak), and glucosephosphate isomerase (Gpi) are all inherited in a Mendelian manner as codominant alleles at nuclear loci. Both parental alleles are equally functional in artificial F₁ hybrids (female B. bombina×male B. variegata) at each of the loci studied. No linkage between any pair of loci was observed. Discovery of this inherited biochemical variation combined with a technique for assaying individual genotypes without killing the animals makes feasible studies of hybrid population structure heretofore impossible.This work was partly supported by the Polish Academy of Sciences within the project MR-II/6.     

Proteomic analysis of muscle between hybrid abalone and parental lines Haliotis gigantea Reeve and Haliotis discus hannai Ino

To understand the potential molecular mechanism of heterosis, protein expression patterns were compared from hybrids of Haliotis gigantea (G) and Haliotis discus hannai (D) using two-dimensional gel electrophoresis (2-DE) and matrix-assisted laser desorption/ionization time-of-flight/time-of-flight analyses. Expression differences were observed in muscle samples from the four groups with 673±21.0 stained spots for H. discus hannai ♀ × H. discus hannai ♂ (DD), 692±25.6 for H. gigantea ♀ × H. gigantea ♂ (GG), 679±16.2 for H. discus hannai ♀ × H. gigantea ♂ (DG) (F₁ hybrid) and 700±19 for H. gigantea ♀ × H. discus hannai ♂ (GD) (F₁ hybrid). Different 2-DE image muscle protein spots had a mirrored relationship between purebreds and the F₁ hybrid, suggesting that all stained spots in F₁ hybrid muscle were on 2-DEs from parents. DD and DG clustered together first, and then clustered with GD, whereas the distance of DD and GG was maximal according to hierarchical cluster analysis. We identified 136 differentially expressed protein spots involved in major biological processes, including energy metabolism and stress response. Most energy metabolism proteins were additive, and stress-induced proteins displayed additivity or over-dominance. In these 136 identified protein spots, hybrid offspring with additivity or over-dominance accounted for 68.38%. Data show that a proteomic approach can provide functional prediction of abalone interspecific hybridization.     

Genetic analysis of heat shock proteins in maize

A genetic analysis of heat shock protein (HSP) synthesis was performed in seedling leaf tissue of two maize inbred lines, their F₁ hybrid and F₂ progeny. Protein synthesis following a high temperature treatment was visualized by [³⁵S]-methionine in vivo labelling and two-dimensional gel electrophoresis. The parental lines' HSP synthesis patterns revealed both qualitative and quantitative polymorphisms implicative of differences in HSP structural genes and regulatory factors. The F₁ hybrid HSP profile indicated that synthesis of all parental HSPs conformed to dominant inheritance patterns, including complete dominance, over-dominance and co-dominance. Alleles for six low-molecularweight HSPs in F₂ progeny assorted according to typical 31 Mendelian ratios for dominant gene expression. There is evidence for unlinked gene loci of four different HSP gene pairs, but data for three other HSP gene pairs were inconclusive, perhaps reflecting linkage for one pair and complex regulatory factor interactions for the other two pairs of genes. These results clearly indicate the existence of genetic variability in HSP synthesis and emphasize the potential of partitioning their roles in thermal tolerance using genetic and molecular analyses.     

A non-additive interaction in a single locus causes a very short root phenotype in wheat

Non-additive allelic interactions underlie over dominant and under dominant inheritance, which explain positive and negative heterosis. These heteroses are often observed in the aboveground traits, but rarely reported in root. We identified a very short root (VSR) phenotype in the F₁ hybrid between the common wheat (Triticum aestivum L.) landrace Chinese Spring and synthetic wheat accession TA4152-71. When germinated in tap water, primary roots of the parental lines reached ~15 cm 10 days after germination, but those of the F₁ hybrid were ~3 cm long. Selfing populations segregated at a 1 (long-root) to 1 (short-root) ratio, indicating that VSR is controlled by a non-additive interaction between two alleles in a single gene locus, designated as Vsr1. Genome mapping localized the Vsr1 locus in a 3.8-cM interval delimited by markers XWL954 and XWL2506 on chromosome arm 5DL. When planted in vermiculite with supplemental fertilizer, the F₁ hybrid had normal root growth, virtually identical to the parental lines, but the advanced backcrossing populations segregated for VSR, indicating that the F₁ VSR expression was suppressed by interactions between other genes in the parental background and the vermiculite conditions. Preliminary physiological analyses showed that the VSR suppression is independent of light status but related to potassium homeostasis. Phenotyping additional hybrids between common wheat and synthetics revealed a high VSR frequency and their segregation data suggested more Vsr loci involved. Because the VSR plants can be regularly maintained and readily phenotyped at the early developmental stage, it provides a model for studies of non-additive interactions in wheat.     

A study on the Changthangi pashmina and the Bakerwali goat breeds in Kashmir: I. Analysis of blood protein polymorphisms and genetic variability within and between the populations

Blood samples of 483 Pashmina goats and 392 Bakerwali goats were taken from the Ladakh and Jammu provinces, respectively, for characterisation of the breeds by polymorphic enzymes and proteins. Furthermore, the sex, age, body weight and hematocrit of both breeds and the pashmina yield of Pashmina goats were recorded. Polymorphisms of 12 enzymes and proteins (albumin (Al), alkaline phosphatase (Alp), amylase (Amy), NADH-diaphorase 1 (Dia1), vitamin-D-binding protein (Gc), haemoglobin (Hb), hemopexin (Hpx), nucleoside phosphorylase (Np), malic enzyme (ME), phosphohexose-isomerase (PHI), transferrin (Tf), X-protein (X)) in blood plasma and hemolysate were determined using gel electrophoresis. Out of 12 protein systems, 10 were found to be polymorphic. In four systems (Al, Amy, Dia1, Hpx) new phenotypes were detected. To estimate the genetic variability within breeds, the degrees of heterozygosity, deviations from the Hardy–Weinberg-equilibrium (HWE) and F_IS-, F_ST-, and F_IT-values were estimated. In comparison with literature data both breeds show slightly higher variability than other goat breeds, with degrees of heterozygosity ranging from 15 to 25% and 24 to 26% in the Pashmina and Bakerwali goat populations, respectively, and the percentage of polymorphic loci ranging from 50 to 67%. On the other hand, a decrease of variability at some loci can be observed in both breeds. Deviations from HWE in combination with a deficiency of heterozygote genotypes were observed in all sub-populations apart from the Pashmina sub-population ‘Likir’. Genetic differences between the goat breeds could be quantified through calculation of Nei’s genetic distances ranging from 0.002 to 0.080. With exception of the Pashmina sub-population ‘Likir’, lower distance values were found between sub-populations within the respective breeds.     

Reproductive and genetic roles of the maternal progenitor in the origin of common wheat (Triticum aestivum L.)

Common wheat (Triticum aestivum L., AABBDD genome) is thought to have emerged through natural hybridization between Triticum turgidum L. (AABB genome) and Aegilops tauschii Coss. (DD genome). Hybridization barriers and doubling of the trihaploid F₁ hybrids’ genome (ABD) via unreduced gamete fusion had key roles in the process. However, how T. turgidum, the maternal progenitor, was involved in these mechanisms remains unknown. An artificial cross‐experiment using 46 cultivated and 31 wild T. turgidum accessions and a single Ae. tauschii tester with a very short genetic distance to the common wheat D genome was conducted. Cytological and quantitative trait locus analyses of F₁ hybrid genome doubling were performed. The crossability and ability to cause hybrid inviability did not greatly differ between the cultivars and wild accessions. The ability to cause hybrid genome doubling was higher in the cultivars. Three novel T. turgidum loci for hybrid genome doubling, which influenced unreduced gamete production in F₁ hybrids, were identified. Cultivated T. turgidum might have increased the probability of the emergence of common wheat through its enhanced ability to cause genome doubling in F₁ hybrids with Ae. tauschii. The ability enhancement might have involved alterations at a relatively small number of loci.     

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.     

Comparative genetic linkage maps of <Emphasis Type="Italic">Eucalyptus grandis</Emphasis>, <Emphasis Type="Italic">Eucalyptus globulus</Emphasis> and their F<Subscript>1</Subscript> hybrid based on a double pseudo-backcross mapping approach

Comparative genetic mapping in interspecific pedigrees presents a powerful approach to study genetic differentiation, genome evolution and reproductive isolation in diverging species. We used this approach for genetic analysis of an F₁ hybrid of two Eucalyptus tree species, Eucalyptus grandis (W. Hill ex Maiden.) and Eucalyptus globulus (Labill.). This wide interspecific cross is characterized by hybrid inviability and hybrid abnormality. Approximately 20% of loci in the genome of the F₁ hybrid are expected to be hemizygous due to a difference in genome size between E. grandis (640 Mbp) and E. globulus (530 Mbp). We investigated the extent of colinearity between the two genomes and the distribution of hemizygous loci in the F₁ hybrid using high-throughput, semi-automated AFLP marker analysis. Two pseudo-backcross families (backcrosses of an F₁ individual to non-parental individuals of the parental species) were each genotyped with more than 800 AFLP markers. This allowed construction of de novo comparative genetic linkage maps of the F₁ hybrid and the two backcross parents. All shared AFLP marker loci in the three single-tree parental maps were found to be colinear and little evidence was found for gross chromosomal rearrangements. Our results suggest that hemizygous AFLP loci are dispersed throughout the E. grandis chromosomes of the F₁ hybrid.Communicated by O. Savolainen     

Genetic evidence for gametophytic selection of wilt resistant alleles in chickpea

Gametophytic selection can drastically reduce the number of selection cycles during crop improvement programs. The objective of the present investigation was to test whether the nature of inheritance of two unlinked disease-resistant loci, h ₁ and h ₂, against Fusarium wilt in chickpea (Cicer arietinum L.) under gametophytic (pollen) selection was similar to that already observed at sporophytic level. A homozygous dominant (H ₁ H ₁ H ₂ H ₂) susceptible genotype JG-62 was crossed to a recessive (h ₁ h ₁ h ₂ h ₂) resistant genotype WR-315 to produce 20 F₁ hybrid seeds. In the following generation, flower buds of 10 F₁ hybrid plants were subjected to toxin stress before anthesis and the remaining ten control F₁ plants’ flowers were sprayed with water. Thirty-four selected BC₁ plants were generated by test crossing resistant WR-315 individuals with pollen from toxin-stressed F₁ individuals. Both control and treated F₁ plants were selfed to produce respective F₂ generations. Two DNA markers, CS-27_700bp and A07C_430bp, linked to susceptible alleles H ₁ and H ₂, respectively, were used to study the inheritance patterns of h ₁ and h ₂ loci in the F₂ and BC₁ generations. One hundred and forty-four selected F₂, 129 control F₂, and 34 selected backcross individuals were tested for the presence or absence of DNA markers. Except for the control F₂, observed ratios of selected F₂ and BC₁ populations exhibited significant chi-square deviations from expected monogenic and digenic ratios. Our results suggest that gametophytic selection is as effective as that realized at the sporophytic level, and that the gametophytic selection can be an effective breeding tool for plant breeding programs.     

Polymorphism and Inheritance of Seed Storage Protein in Sunflower

The data on polymorphism and inheritance of the seed storage protein helianthinin are presented. The results of hybrid analysis indicate that in the annual sunflower Helianthus annuus, helianthinin synthesis is controlled by at least three loci: HelA, HelB, and HelC. Codominant alleles controlling different electrophoretic variants of polypeptides were identified at each of the loci. The HelA locus was inherited independently of HelB and HelC in a series of dihybrid crosses. The frequencies of recombination between loci HelB and HelC estimated in F₂ and BC of two crossing combinations were respectively 21.8 and 19.0%. Segregation of the HelC-controlled variants in the progenies from the crosses of cultured sunflower with annual wild species and forms corresponded to that theoretically expected for Mendelian inheritance. The maternal type of helianthinin inheritance was observed in the progenies from the crosses of inbred H. annuus L. lines with perennial diploid and polyploid HelianthusL. species. Altered expression of the HelClocus was detected in some hybrid combinations. These alterations appeared in early (F₁, F₂) hybrid generations and were similar in different hybrid combinations. They did not depend on the perennial paternal species being more influenced by the maternal genotype and by the mode of obtaining hybrids (in an embryo culture or in the field). These results are explained by genomic shock generated by hybridization of genetically incompatible species.     

Defensive behaviour and its inheritance in the anabantoid fish, Macropodus opercularis and Macropodus opercularis concolor

In this preliminary study defense behaviour patterns (fear responses) are described in two closely related, behaviourally different inbred labyrinth fish subspecies and in their F₁ generation. The subspecies M. opercularis (characterized briefly by “active escape”) and M. opercularis concolor (characterized by “passive escape”) showed specific differences in the manifestation of certain defense behaviour patterns. In the F₁ hybrid generation dominance and overdominance of M. opercularis was found in most defense behaviour patterns. Analysing the frequencies and sequences of movement patterns it could be shown that defensive behaviour is not a random or entirely “plastic” process but that there is sequential linkage between the patterns and they form characteristic clusters. Our results suggest that manifestations of different patterns are under genetic control and presumably, genetic determination of certain patterns is not very complex. Attempts were made to determine whole brain noradrenaline, serotonine and dopamine levels of the two subspecies and a significant difference was found in the noradrenaline content.     

Constructing genetic linkage maps for Pinus elliottii var. elliottii and Pinus caribaea var. hondurensis using SRAP, SSR, EST and ISSR markers

A total of 122 F₁ individuals from a single full-sib interspecific hybrid family crossed between Pinus elliottii var. elliottii (PEE) and P. caribaea var. hondurensis (PCH) were used to construct a detailed genetic linkage maps using four types of molecular markers: sequence-related amplified polymorphism (SRAP), microsatellite (SSR), expressed sequence tag polymorphism (ESTP) and inter-simple sequence repeat (ISSR). There were 381 SRAP, 108 SSR, 25 ESTP and 32 ISSR loci, segregating in the interspecific F₁ hybrid individuals. Framework maps were constructed at a LOD score threshold of 4.0 using the JoinMap^® 3.0. The map for the male parent (PCH) had 108 markers in 16 linkage groups (LGs), with a total length of 1,065.9 cM (Kosambi) and an average marker interval of 9.87 cM. The map for the female parent (PEE) contained 93 markers in 19 LGs, with a total length of 1,006.7 cM (Kosambi) and an average marker interval of 10.82 cM. The maps for PCH and PEE covered 56.5 and 70.3 % of their respective genomes. Based on the position of 36 loci segregating in both parents, 8 homologous LGs between PEE and PCH were identified.