Detection and inheritance of RFLPs in Eucalyptus nitens

The level of polymorphism using genomic and cDNA probes with a number of restriction enzymes and the inheritance of the RFLP loci was investigated in E. nitens. The polymorphism detected with 366 genomic and cDNA probes and three to six restriction enzymes was analysed in three-generation outbred pedigrees. No difference in the level of polymorphism detected with genomic versus cDNA probes was observed. There was a difference in the efficiency of detection of polymorphism with six different restriction enzymes, with three of the enzymes (BglII, DraI and EcoRI) showing substantially more polymorphism than the others. There was no significant correlation between the size of the DNA fragments generated by the enzymes and the detection of polymorphism. Several cases of restriction-site mutations resulting in a polymorphism were observed. The inheritance of 69 loci was analysed in two pedigrees resulting from interpopulational crosses. The majority of the loci segregated according to expected ratios with distortion observed in only 3% of loci. Probes from the cDNA library detected a greater proportion of loci with more than two alleles than did probes from the genomic library. The high polymorphism, large number of alleles, and ease of interpretation of RFLPs in E. nitens means that they will be useful in a range of applications such as genetic linkage maps and paternity analysis.     

Genetic parameters for growth, wood density and pulp yield in Eucalyptus globulus

Genetic variation and co-variation among the key pulpwood selection traits for Eucalyptus globulus were estimated for a range of sites in Portugal, with the aim of improving genetic parameters used to predict breeding values and correlated response to selection. The trials comprised clonally replicated full-sib families (eight trials) and unrelated clones (17 trials), and exhibited varying levels of pedigree connectivity. The traits studied were stem diameter at breast height, Pilodyn penetration (an indirect measure of wood basic density) and near infrared reflectance predicted pulp yield. Univariate and multivariate linear mixed models were fitted within and across sites, and estimates of additive genetic, total genetic, environmental and phenotypic variances and covariances were obtained. All traits studied exhibited significant levels of additive genetic variation. The average estimated within-site narrow-sense heritability was 0.19 ± 0.03 for diameter and 0.29 ± 0.03 for Pilodyn penetration, and the pooled estimate for predicted pulp yield was 0.42 ± 0.14. When they could be tested, dominance and epistatic effects were generally not statistically significant, although broad-sense heritability estimates were slightly higher than narrow-sense heritability estimates. Averaged across trials, positive additive (0.64 ± 0.08), total genetic (0.58 ± 0.04), environmental (0.38 ± 0.03) and phenotypic (0.43 ± 0.02) correlation estimates were consistently obtained between diameter and Pilodyn penetration. This data argues for at least some form of pleiotropic relationship between these two traits and that selection for fast growth will adversely affect wood density in this population. Estimates of the across-site genetic correlations for diameter and Pilodyn penetration were high, indicating that the genotype by environment interaction is low across the range of sites tested. This result supports the use of single aggregated selection criteria for growth and wood density across planting environments in Portugal, as opposed to having to select for performance in different environments.     

Photosynthetic responses to overnight frost in Eucalyptus nitens and E. globulus

Significant expansion in the area of eucalypt plantations in Tasmania has led to their establishment at altitudes that are close to the upper limits of the planting distributions of Eucalyptus nitens and E. globulus, the main species planted. This has implications for plantation productivity. We investigated the processes that limit productivity in these environments through a study of freezing-induced depression of photosynthesis of E. nitens saplings in the field and plantlets of E. nitens and E. globulus clones in a controlled environment cabinet. In the field consecutive frosts of around –4.6°C had a cumulative effect, reducing maximum net photosynthesis ( A _max) by 17%, and then a further 9%, respectively, compared with saplings insulated from the frosts. Shading saplings pre-dawn had no effect on A _max measured after 1030 hours indicating that the reduction in A _max at this time was independent of photoinhibition. Recovery of A _max to pre-frost levels required at least two consecutive frost-free nights and was dependent on the severity of frost. Photosynthetic light response curves indicated that reduced A _max was associated also with decreased quantum yield and stomatal conductance. Similar intracellular carbon dioxide concentration between exposed and insulated saplings indicated that low stomatal conductance did not limit photosynthesis through carbon dioxide limitation. The timing of frost events was critical: E. nitens saplings took less time to recover from reduced A _max in the field when they were hardened. Unhardened plantlets of E. nitens and E. globulus clones had greater reduction of A _max and took longer to recover from frost events than hardened plantlets. E. globulus was more susceptible to frost-induced reduction of A _max than E. nitens. This is consistent with its planting range which is restricted to mild sites compared with that of E. nitens.     

Mapping of quantitative trait loci influencing frost tolerance in Eucalyptus nitens

 Regions of the genome influencing frost tolerance in an outbred family of Eucalyptus nitens have been identified. Two QTLs present on the same linkage group, but located 40 cM apart, were identified using single-factor analysis of variance. The QTLs explained between 7.7 and 10.8% of the phenotypic variation for frost tolerance in this family. Analysis of marker loci linked to the QTLs showed one of them to have a simple mode of action with the effect segregating from the male parent in the family. For the other QTL multiple alleles were identified. This QTL showed segregation from the female parent which gave a positive effect on frost tolerance; however, an allele segregating from the male parent was identified which showed a negative interaction with the allele for increased frost tolerance. Received: 10 May 1997 / Accepted: 2 June 1997     

Genetic variation in Eucalyptus nitens pulpwood and wood shrinkage traits

Eucalyptus nitens plantations are generally established for pulpwood production but an increasing area is being managed for solid wood. Genetic variation in, and correlations among, three Kraft pulpwood traits (diameter at breast height, basic density and near-infrared-predicted cellulose content) and three 12-mm wood-core shrinkage traits (recoverable collapse, net shrinkage and gross shrinkage) were examined, utilising data from two 9-year-old first-generation progeny trials in Tasmania. These trials contained approximately 400 open-pollinated families (over 100 of which were sampled for wood properties) representing three central-Victorian E. nitens races. Significant genetic variation at the race and/or within-race level was identified in all traits. Within races, relative levels of additive genetic variation were higher for shrinkage traits, although narrow-sense heritabilities were lower and the expression of genetic variation less stable across sites than for other wood property traits. Heterogeneous intertrait genetic correlations were identified across sites between growth and some wood property traits. However, where significant, genetic correlations indicated that within-race selection for growth would adversely affect core basic density and all core shrinkage traits. Furthermore, results based on cores suggested that within-race selection for higher basic density would favourably impact on cellulose content and collapse but selection for either higher basic density or cellulose content would adversely affect net shrinkage. Most within-race genetic variation in gross shrinkage appeared to be due to genetic variation in collapse. The implications of these results for sawn timber breeding will depend on the strength of genetic correlations between core traits and rotation-age objective traits and objective trait economic weights.     

Genetic causes of heterosis in juvenile aspen:a quantitative comparison across intra- and inter-specific hybrids

The genetic causes of heterosis in tree growth were investigated by a comparative genetic analysis of intra- and inter-specific crosses derived from Populus tremuloides and P. tremula. A new analytical method was developed to estimate the effective number of loci affecting a quantitative trait and the magnitudes of their additive and dominant effects across loci. The method combines the assumption of multiple alleles, as frequently found in outcrossing species, and the family structure analysis at different hierarchical levels. During the first 3 years of growth, interspecific hybrids displayed strong heterosis in stem growth, especially volume index, over intraspecific hybrids. By a series of joint analyses on the combining ability and the genetic component, we found that F₁ heterosis might be due to overdominant interaction between two alleles, one from the P. tremuloides parent and the other from the P. tremula parent, at the same loci. This inference was derived from the finding that heterozygotes, newly formed through species combination, showed much greater growth than the heterozygotes from intraspecifc crosses at a reference locus. Heterosis in aspen growth appeared to be under multi-genic control, with a slightly larger number of loci for stem diameter and volume (9–10) than for height (6–8). For traits with non-significant heterosis, such as stem allometry and internode number and length, the number of underlying loci seemed to be much fewer (3–4). While additive effects appeared to influence seedling traits collectively across loci, a few major dominant loci had much larger effects on stem growth.     

Chilling and freezing stress in live oaks (Quercus section Virentes): intra- and inter-specific variation in PS II sensitivity corresponds to latitude of origin

Sensitivity to cold and freezing differs between populations within two species of live oaks (Quercus section Virentes Nixon) corresponding to the climates from which they originate. Two populations of Quercus virginiana (originating from North Carolina and north central Florida) and two populations of the sister species, Q. oleoides, (originating from Belize and Costa Rica) were grown under controlled climate regimes simulating tropical and temperate conditions. Three experiments were conducted in order to test for differentiation in cold and freezing tolerance between the two species and between the two populations within each species. In the first experiment, divergences in response to cold were tested for by examining photosystem II (PS II) photosynthetic yield (ΔF/F _m′) and non-photochemical quenching (NPQ) of plants in both growing conditions after short-term exposure to three temperatures (6, 15 and 30°C) under moderate light (400 μmol m⁻² s⁻¹). Without cold acclimation (tropical treatment), the North Carolina population showed the highest photosynthetic yield in response to chilling temperatures (6°C). Both ecotypes of both species showed maximum ΔF/F _m′ and minimum NPQ at their daytime growth temperatures (30°C and 15°C for the tropical and temperate treatments, respectively). Under the temperate treatment where plants were allowed to acclimate to cold, the Q. virginiana populations showed greater NPQ under chilling temperatures than Q. oleoides populations, suggesting enhanced mechanisms of photoprotective energy dissipation in the more temperate species. In the second and third experiments, inter- and intra-specific differentiation in response to freezing was tested for by examining dark-adapted F _v/F _m before and after overnight freezing cycles. Without cold acclimation, the extent of post-freezing declines in F _v/F _m were dependent on the minimum freezing temperature (0, −2, −5 or −10°C) for both populations in both species. The most marked declines in F _v/F _m occurred after freezing at −10°C, measured 24 h after freezing. These declines were continuous and irreversible over the time period. The North Carolina population, however, which represents the northern range limit of Q. virginiana, showed significantly less decline in F _v/F _m than the north central Florida population, which in turn showed a lower decline in Fv/F _m than the two Q. oleoides populations from Belize and Costa Rica. In contrast, after exposure to three months of chilling temperatures (temperate treatment), the two Q. virginiana populations showed no decline in F _v/F _m after freezing at −10°C, while the two Q. oleoides populations showed declines in F _v/F _m reaching 0.2 and 0.1 for Costa Rica and Belize, respectively. Under warm growth conditions, the two species showed different F ₀ dynamics directly after freezing. The two Q. oleoides populations showed an initial rise in F ₀ 30 min after freezing, followed by a subsequent decrease, while the Q. virginiana populations showed a continuous decrease in F ₀ after freezing. The North Carolina population of Q. virginiana showed a tendency toward deciduousness in response to winter temperatures, dropping 58% of its leaves over the three month winter period compared to only 6% in the tropical treatment. In contrast, the Florida population dropped 38% of its leaves during winter. The two populations of the tropical Q. oleoides showed no change in leaf drop during the 3-months winter (10% and 12%) relative to their leaf drop over the same timecourse in the tropical treatment. These results indicate important ecotypic differences in sensitivity to freezing and cold stress between the two populations of Q. virginiana as well as between the two species, corresponding to their climates of origin.     

Resistance of tomato F1 hybrids to grey mold

A two-year estimation of Botrytis cinerea stem damage of 299 experimental F₁hybrids grown in 2 rotations: winter-autumn and summer-autumn was carried out. The estimation was made in a period of mass fruitage. As a result it was shown, that the hybrids with determinate type of growth (sp) are damaged more strongly than indeterminate ones (sp +). An average disease severity in the first group was 2.8 against 2.3 in the second one using a scale 0–4. Number of strongly injured (2.5–4) plants in each group were 44.1 % and 30.7 % and number of moderate resistant (0–1.5 numbers) plants were 10 % and 8.5 % accordingly. The analysis of resistance of F₁ hybrids determined significant influence of parental lines on a degree of a damage of plant stems by B. cinerea.     

AFLP genetic maps of Eucalyptus globulus and E. tereticornis

 Amplified fragment length polymorphism (AFLP) analysis is a rapid and efficient technique for detecting large numbers of DNA markers in eucalypts. We have used AFLP markers in a two-way pseudo-testcross strategy to generate genetic maps of two clones of different Eucalyptus species (E. tereticornis and E. globulus). Of 606 polymorphic fragments scored, 487 segregated in a 1 : 1 ratio, corresponding to DNA polymorphisms heterozygous in one parent and null in the other. In the maternal E. tereticornis map, 268 markers were ordered in 14 linkage groups (919 cM); the paternal E. globulus map had 200 markers in 16 linkage groups (967 cM). Results from PGRI software were compared with MAPMAKER. The average density of markers was approximately 1 per 3.9 cM. Framework markers were ordered with an average confidence level of 90%, covering 80–100% of the estimated Eucalyptus genome size. In order to investigate the homologies between the E. tereticornis and the E. globulus genetic linkage maps, we included 19 markers segregating 3 : 1 in the analysis. Some homeologous linkage groups were recognized. The linkage data developed in these maps will be used to detect loci controlling commercially important traits. Received: 17 July 1997 / Accepted: 13 October 1997     

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.     

Identification and mode of action of quantitative trait loci affecting seedling height and leaf area in Eucalyptus nitens

 Regions of the genome influencing height and leaf area in seedlings of a three-generation outbred pedigree of Eucalyptus nitens have been identified. Three QTLs affecting height and two QTLs affecting leaf area were located using single-factor analysis of variance. The three QTLs affecting height each explained between 10.3 and 14.7% of the phenotypic variance, while the two QTLs for leaf area each explained between 9.8 and 11.6% of the phenotypic variation. Analysis of fully informative marker loci linked to the QTLs enabled the mode of action of the QTLs to be investigated. For three loci the QTL effect segregated from only one parent, while for two loci the QTL showed multiple alleles and the effect segregated from both parents in the pedigree. The two QTLs affecting leaf area were located in the same regions as two of the QTLs affecting height. Analysis of these regions with fully informative markers showed that both QTLs were linked to the same markers, but one had a similar size of effects and a similar mode of action for both height and leaf area, whilst the other showed a different mode of action for the two traits. These regions may contain two closely linked genes or may involve a single gene with a pleiotrophic effect on both height and leaf area. The QTL with the greatest effect showed multiple alleles and an intra-locus interaction that reduced the size of the effect. Assessment for two of the QTLs in a second related family did not show an effect associated with the marker loci; however, this was consistent with the mode of action of these QTLs and the pattern of inheritance in the second family. Received: 1 August 1996 / Accepted: 25 October 1996     

Daily stem growth patterns in irrigated Eucalyptus globulus and E. nitens in relation to climate

Daily increments in stem radius were determined from hourly dendrometer measurements in each of three irrigated Eucalyptus nitens and E. globulus trees. Multiple regressions determined from daily weather variables accounted for 40–50% of the variance in increment. The use of weather variables lagged by 1–2 days increased the variance explained. The diurnal variation in stem radius was resolved into three mathematically defined phases: shrinkage, recovery and increment. The positive daily net increment in stem radius, by definition, occurred in the increment phase. Average weather conditions during this phase (predominantly night-time) did not explain any more variance in increment than the average daily conditions, determined over a 24 h period. Daily increment was resolved into a rate of stem radius increase during the increment phase and the duration (hours) of that phase. Significant species by month interactions were evident with growth in summer characterised by faster rates of stem expansion over shorter time periods within each diurnal cycle. E. nitens tended to have longer increment phases in spring and autumn, and faster phase rates in autumn than E. globulus. Interactions between weather variables and cambial growth were complicated and varied over the year. The correlation between temperature and stem growth varied from positive in spring to zero or negative during summer. The data indicate a need to understand weather-by- climate interactions at the level of whole tree physiology in order to fully understand the effect of weather on cambial activity and therefore stem increment and wood properties. Received: 12 April 1999 / Accepted: 6 July 1999     

Multivariate analysis of tooth crown morphology in Japanese-American F1 hybrids

Mesiodistal crown diameters of I1 through M1 and six non-metric crown traits in permanent dentition of Japanese-American F₁ hybrids were compared with those of the parental populations. The hybrids were born of Japanese females and American males, both Caucasians and American Blacks, after World War II and brought up at Elizabeth Saunders Home in Kanagawa Prefecture, Japan. The comparisons were undertaken by means of multivariate analysis methods such as principal component analysis, distance and similarity coefficients and multidimensional scaling. The F₁ hybrids generally occupy an intermediate position of the two parental populations, and this is particularly evident in size component of the crown measurements and in distance analysis of frequencies of the non-metric crown traits. The shape component of the crown measurements, however, not necessarily follows such a rule and suggests a more complicated gene control than in the case of the size component. Also, each measurement and frequency of non-metric trait of tooth crowns in F₁ hybrids is not uniformly at the middle position between their parental populations but this is true when they are analyzed as multivariables.     

Genetic dissection of vegetative propagation traits in Eucalyptus tereticornis and E. globulus

We have detected quantitative trait loci (QTLs) affecting vegetative propagation traits in Eucalyptus tereticornis and Eucalyptus globulus. Using amplified fragment length polymorphism (AFLP) genetic linkage maps, the inheritance of 199 markers was assessed in 94 F₁ individuals with extreme adventitious rooting response, and in 221 randomly chosen F₁ individuals. Phenotypes were scored in 1995 and 1996. QTL analyses were performed using chi-square tests (χ²), single-marker analysis (SMA), interval mapping (IM) and composite interval mapping (CIM). All approaches yielded similar QTL detection results. Three QTLs are hypothesized for mortality (MORT=% dead cuttings), nine for adventitious rooting (ROOT, RCT=% rooted cuttings relative to the surviving or total cuttings, respectively), four for petrification (PETR=% surviving unrooted cuttings), one for sprouting ability (SPR=number of stump sprout cuttings harvested in 1995) and four for the stability of adventitious rooting (STAB=absolute value of the difference ROOT95-ROOT96). All putative QTLs for MORT and PETR were located on the E. tereticornis map, and for SPR and STAB on the E. globulus map. We found different QTLs for MORT, ROOT, RCT, SPR and STAB. Putative QTLs accounted for 2.6–17.0% of the phenotypic variance of a trait (R²). Estimated standardized gene substitution effects varied between 0.13 and 0.49 phenotypic standard deviations (σ_p). These results indicate that the phenotypic variation in these traits has a meaningful genetic component and that stable QTLs can be found in a family of reasonable size where no previous knowledge of the trait was available. Received: 1 September 1998 / Accepted: 24 February 1999     

Cold hardening reduces photoinhibition of Eucalypts nitens and E. pauciflora at frost temperatures

Photoinhibition of photosynthesis at low temperatures was investigated in two species of subalpine eucalypt, Eucalypts nitens (Deane and Maiden) Maiden and E. pauciflora Sieb. ex Spreng. Imposition of an artificial cold-hardening treatment increased the frost tolerance of leaf tissue and increased tolerance to excess light. Cold-hardened seedlings of both species had a higher photosynthetic capacity than non-hardened seedlings at 6 and 16°C and lower levels of non-photochemical quenching (NPQ) at 20 and 5°C. Furthermore, hardened seedlings had faster rates of NPQ development at 5 and −3.5°C. An increase in minimal fluorescence, which indicates slowly reversible photoinhibition, was evident in all seedlings at −1.5 and −3.5°C but was less pronounced in hardened seedlings, with a threefold faster rate of development of NPQ, at −3.5°C than non-hardened seedlings. Hardened seedlings also recovered faster from photoinhibition at −3.5°C. Thus cold hardening increased tolerance to high light in these species. Differences between E. nitens and E. pauciflora in their response to excess light were small and significant only at −3.5°C. Faster recovery from photoinhibition of E. pauciflora was consistent with its occurrence in colder habitats than E. nitens. Received: 27 April 1997 / Accepted: 9 September 1997     

Natural hybridization between an outcrossing and a selfingPhlox (Polemoniaceae): The maternal species of F1 hybrids

Natural F₁ hybrids between the outcrossingPhlox drummondii and the predominantly selfingP. cuspidata were examined to ascertain the proportion of hybrid individuals mothered by each species. Species-specific restriction fragment patterns (both nrDNA and cpDNA) were established as markers, and synthetic hybrids of known parentage were utilized to determine that the chloroplast genome is maternally inherited. Of 89 mature natural hybrids examined, approximately two thirds were mothered byP. drummondii, the outcrosser. That the outcrosser should mother most hybrids is expected since it is dependent upon incoming pollen for fertilization, and hybrids may result when heterospecific pollen is received. The fact that the highly selfingP. cuspidata mothered nearly one third of the hybrids is surprising, and may be related to both pre-zygotic and post-zygotic factors. Which species mothers hybrids has important implications for the potential for introgression as well as its direction.     

Root and shoot elongation of rhizotron-grown seedlings of Eucalyptus nitens and Eucalyptus globulus in relation to temperature

Information on the growth response of a crop plant in relation to temperature can be helpful in selecting genotypes to suit local environments, scheduling favourable time of planting and forecasting growth and yield. To determine the effects of varying temperature on root and shoot elongation of eucalypt seedlings, elongation rates of roots and shoots were measured in rhizotrons for two species (Eucalyptus nitens (Deane and Maiden) Maiden, and Eucalyptus globulus Labill.) at a temperature range of 5–23 °C. Within this range of temperatures, elongation rates of roots and shoots of both species increased with an increase in temperature. Roots of E. globulus were more sensitive and shoots less sensitive to temperature than those of E. nitens. However, the threshold temperature corresponding with zero elongation rate predicted from the regression of elongation rate against temperature was similar for the roots (∼5 °C) and shoots (∼0 °C) of both species. Hysteresis did not appear to have a significant influence on root or shoot elongation of both species during warming compared with cooling. Results are discussed highlighting the importance of the interaction between development and growth of plant components.     

Genetic structures of allopatric and sympatric populations in Pseudorasbora pumila pumila and Pseudorasbora parva

Pseudorasbora pumila pumila was collected to investigate its genetic relationships from 11 localities in Hokkaido and eastern Honshu of Japan with Pseudorasbora parva from 6 localities in Hokkaido, eastern Honshu, and Polovynka River (a tributary of the Amur River in eastern Russia) in 1996–1998. In Hokkaido and Akita prefectures (Honshu), three P. pumila pumila populations were distributed with P. parva sympatrically. Samples of P. pumila pumila and P. parva were analyzed based on 27 allozyme loci. Genetic differentiation between P. pumila pumila and P. parva (D = 0.522–0.622, average 0.601) was almost equal to or greater than that between other congeneric fish species previously reported. In the three sympatric populations of two species, which had arisen from artificial transplantation of the latter into localities of the former, the individual frequency composition of genetic types (P. pumila pumila, P. parva, F₁ hybrid, and later filial generation hybrid) were significantly different from each other. Differing from the previous studies, F₁ hybrids were fertile and coexisting populations of the two species included the F₁ hybrid and the later filial generation hybrid. These results suggest that genetic population dynamics of sympatric populations of the two species would be influenced not only by hybridization but also by ecological and environmental factors. Alternatively, the existence of populations that possessed both the alleles specific to P. pumila pumila and those specific to P. parva may indicate that introgressive hybridization, instead of species displacement, has occurred between the two species under a certain condition.     

Genetic segregation in relation to chromosome pairing in tetraploid hybrids between Lolium perenne and L. multiflorum

Summary Segregation at one of the loci controlling tiller-base pigmentation was studied to determine the mode of inheritance in tetraploid hybrids between Lolium perenne and L. multiflorum. The results could be explained by tetrasomic inheritance and thus did not support previous reports of a degree of preferential chromosome pairing in this material. However, double reduction and aneuploidy may to some extent have masked any tendency to disomic segregation brought about by preferential pairing. Moreover, there was significant heterogeneity between families in the segregation ratios which may indicate genetically controlled differences in pairing behaviour. The results are related to previous cytological and genetic studies.