Genetic parameters and provenance variation of Pinus radiata D. Don. ‘Eldridge collection’ in Australia 1: growth and form traits

Growth and form traits data were obtained from eight provenance trials of radiata pine (Pinus radiata D. Don) planted across the radiata pine plantation estate in southeast Australia. The genetic pool included 466 open-pollinated families collected from A?o Nuevo, Monterey and Cambria provenances on the Californian mainland coast in the USA and from Guadalupe and Cedros islands off the coast of Baja California in Mexico. Early survival of all provenances was around 90%, except for Cedros (<60%). Monterey and A?o Nuevo were the best performers at almost all sites. However, good growth performance of Cambria and good stem straightness of Guadalupe on some sites are important results, because the genetic base of the present Australian plantations evidently originated from only Monterey and A?o Nuevo. The average estimated single-site heritability for diameter at breast height was 0.22 and 0.32 at juvenile and mature ages, respectively. Heritability estimates for stem straightness and branching ranged from 0.23 to 0.55. Genetic correlation estimates between diameter at breast height (DBH) at juvenile and rotation ages were all >0.80. Estimates of between-site genetic and provenance correlations for DBH were often low, indicating high genotype by environment interaction across trials, consistent with previous Australian studies. However, there was minimal G × E among trials on high-altitude high-rainfall sites and among trials on low-altitude, low-rainfall sites.     

Genetic parameters and provenance variation of Pinus radiata D. Don. ‘Eldridge collection’ in Australia 2: wood properties

Provenance variation and genetic parameters for wood properties of mature radiata pine (Pinus radiata D. Don) were studied by sampling three provenance/progeny trials in southeast Australia. Among the mainland provenances, Monterey and Año Nuevo had higher density and modulus of elasticity (at one site) than Cambria. Basic density and predicted modulus of elasticity (MoE) for the island provenances, Guadalupe and Cedros, were ～20% higher at Billapaloola compared to mainland provenances grown at Green Hills and Salicki, differences that may or may not be linked to site differences. Heritability estimates of density, predicted MoE and microfibril angle were significant and $ {\bar{h}^2} $ ?>?0.45, suggesting moderate to strong genetic control. The estimated genetic correlations between diameter at breast height and wood properties in the current study were weaker (less negative) than the mean estimated from the current breeding population generation in radiata pine. Of the wood properties, density showed the strongest adverse genetic correlations with growth (mean r _A ?=??0.23?±?0.09). Selection for MoE may produce greater gain than selection for density because MoE had almost twice the estimated additive genetic coefficient of variation ( $ {\overline {\text{CV}}_A} $ ) compared to density. Estimated type B genetic correlations (r _B) for all wood quality traits were typically high, conforming to the trend that wood properties have low genotype-by-environment interaction (G?×?E). Significant differences in wood properties among provenances, families and/or individual trees provide an opportunity for breeding programmes to select superior trees for solid wood production that will combine superior growth with desirable wood traits.     

Genetic stability of wood density and diameter in <Emphasis Type="Italic">Pinus radiata</Emphasis> D. Don plantation estate across Australia

Genetic variation for wood quality traits and diameter growth for radiata pine (Pinus radiata D. Don) at age 20/21 years was estimated from eight trials in Australia. The traits studied were wood density, acoustic time-of-flight (an indirect measure of stiffness) and diameter at breast height (DBH). Wood density and DBH exhibited significant additive genetic variation whereas non-additive effects were not significantly different from zero. Time of flight was also not significantly different from zero for both additive and non-additive effects, respectively. Average single-site heritability estimates (±SE) for wood density and DBH were 0.38 ± 0.10 and 0.16 ± 0.08, respectively. Pooled-site heritability estimates for wood density and DBH were 0.38 ± 0.10 and 0.08 ± 0.10, respectively. For density, there was little evidence of genotype-by-environment interaction (GEI) across the eight trials at the additive level (type B additive genetic correlation; r _BADD = 0.73 ± 0.08) and type B genetic correlation for full-sib families (r _BFS = 0.64 ± 0.08). In contrast, the type B additive genetic correlation for DBH was lower, (r _BADD = 0.51 ± 0.14), suggesting evidence of GEI. However, type B genetic correlation for full-sib families was moderate (0.63 ± 0.11) for DBH, suggesting that there may be some stable full-sib families. On the basis of the results of this study, GEI should be considered in order to optimise deployment of improved germplasm in Australia.     

红锥家系遗传变异与优良家系选择

为选择红锥(Castanopsis hystrix)优良家系,对2–11年生红锥家系的生长性状和遗传参数进行了研究分析。结果表明,家系间在树高、胸径和单株材积上的差异达显著或极显著水平,家系的树高、胸径和单株材积受中等或中等以上遗传控制,具有很强的家系和家系内单株选择潜力;家系的单株材积变异幅度较大,胸径居中,树高的较小,显示出红锥家系间存在着丰富的变异;家系和单株遗传力随林龄的增大逐渐变小,到7–9年生时逐渐稳定。年度生长性状的相关分析表明,树高、胸径和单株材积的遗传相关系数随林龄逐渐增大,到7年生时最大,分别达到0.9602、0.9340和0.9849,之后趋于稳定,因此,7年生可作为早期选择的适合年龄。结合早期选择及形质指标,11年生时最终选择出优良家系10个,其平均树高、胸径和单株材积分别为13.95 m、14.34 cm和0.1229 m3,平均遗传增益分别为12.24%、18.69%和51.59%,其通直度、圆满度、分枝角度、最大分枝、冠幅、枝下高等形质指标也提高7.71%~12.94%。     

Genotype by environment interactions for Pinus radiata in New South Wales, Australia

Historical data from 26 progeny trials in the NSW breeding program was analyzed to determine the extent and practical importance of genotype by environment (G×E) interactions for a range of traits. Significant G×E interaction was present for diameter (DBH) with 75% of the 35 estimated between-site genetic correlations falling below the threshold value of 0.6 where regionalization should be considered. Estimated genetic correlations for stem straightness, branch quality, and outerwood density were much higher, indicating these traits are not as affected by G×E. Levels of G×E for DBH are sufficient to have serious impacts on the expression of genetic gain in deployed material. For DBH, altitude differences between sites appear to be the key factor driving the G×E with a difference of greater than 280 m in altitude leading to a breakdown in correlations. Two groups of sites were identified as having limited G×E within each group: one for sites above 900 m elevation plus a lower-altitude group. Sites included in the higher-altitude group were located across the entire north–south geographic range of NSW. Equations for prediction of site mean DBH indicate that altitude, prior land use, and underlying geology are key driving variables. A more complex model was developed for predicting the between-site genetic correlations for DBH with the model accounting for approximately 50% of the observed variation.     

The origin and genetic diversity of Pinus radiata in Australia

Summary Despite the fact that forest trees are in early stages of domestication there has been little direct evaluation of either the origin of, or genetic diversity within the breeding material in tree improvement programs. Allozyme variation was used to compare the total genetic diversity in the breeding programs of P. radiata within Australia and the five wild populations in North America. The current breeding populations were very similar genetically and were essentially homogenous with only 1.8% of the variation among programs. The total genetic diversity in the species was 0.12, which is a low estimate compared to most conifers. Overall in the Australian material the genetic diversity was somewhat less. The comparison of allelic frequencies in the five native populations with the Australian material indicates that the Monterey and Año Nuevo populations were probably the major source of the original introductions and that a substantial portion of the genetic diversity in the two populations has been captured in current breeding programs. The three southern populations do not appear to be currently represented in the breeding programs. The implications for future breeding strategies are discussed.     

Predicted genetic gain and realised gain in stand volume of Eucalyptus globulus

Realised gains in stand volume of Eucalyptus globulus Labill. families at 7–9 years were compared with gains predicted at 3.5 years. Gain predictions were based on height, diameter at breast height (DBH), sectional area, and stem volume for 153 full-sib families and 18 commercial checks in five-tree line plots on three West Australian sites. Single-site narrow-sense heritability estimates were 0.12–0.24 for height, 0.08–0.12 for DBH, 0.09–0.13 for sectional area, and 0.14–0.19 for stem volume. Genetic dominance effects were significant (p?<?0.05) in most cases, and the estimated dominance/additive genetic variance ratio was heterogeneous for height and DBH. Stand volume was measured for 93 of the same families and checks in 40-tree block plots on four sites. Heritability of stand volume was 0.25–0.76, with an across-sites estimate of 0.41. Dominance effects were statistically absent at two sites. Estimated region-wide additive genetic correlations between selection traits (in line plots) and stand volume (in block plots) ranged from 0.86 to 0.90. Estimated stand volume gain was 23 % of the mean for the best 12 % of families and 14 % of the mean for the best 24 % of families. Realised gain was under-estimated by predictions based on height, DBH, and sectional area, which had smaller coefficients of additive variation than did stem volume. It is concluded that although BLUP analysis of early-age height and DBH can provide for indirect selection on E. globulus stand volume, analysis of stem volume is required to predict genetic gain at an appropriate scale.     

Genetic parameter estimates in a clonally replicated progeny test of teak (Tectona grandis Linn. f.)

Teak (Tectona grandis Linn. f.) has been planted extensively in the tropics for its highly valued timber. We analysed data from a 3.5-year-old teak progeny test with clonal replication located in northern Australia. Additive and non-additive genetic variances were estimated for commercially important traits. Trees originating from seedlings were on average 2% taller and 4% straighter than those of the same genotype originating from cuttings. Non-additive genetic variance represented 35–50% of total genetic variance for growth traits and 63% of total genetic variance for incidence of flowering. Narrow-sense heritability was 0.22 for diameter, 0.18 for height and volume, 0.07 for stem straightness, 0.05 for insect defoliation, 0.03 for epicormic sprouts and 0.30 for incidence of flowering (estimated on an assumed underlying continuous scale). Broad-sense heritability was 0.37 for diameter, 0.28 for height, 0.35 for volume, 0.12 for stem straightness, 0.06 for insect defoliation, 0.12 for epicormic sprouts and 0.71 for incidence of flowering. Positive correlations were found between tree volume and flowering and between tree volume and stem straightness. The presence of sizeable non-additive variance supports the selection and deployment of clones to capture the full extent of genetic variation in commercially important traits.     

Genetic variation for growth,morphological, and physiological traits in a wild population of the Neotropical shade­tolerant rainforest tree <Emphasis Type="BoldItalic">Sextonia rubra</Emphasis> (Mez) van der Werff (Lauraceae)

Quantitative genetic diversity is a fundamental component of the interaction between natural populations and their environment. In breeding programmes, quantitative genetic studies on tropical trees have so far focused on fast-growing, light-demanding species, but no information exists on shade-tolerant, slow-growing species. For this study, 27 3-year-old open-pollinated families of the Neotropical shade-tolerant rainforest tree Sextonia rubra were measured in semicontrolled conditions for 20 morphological, growth, and photosynthesis traits; the effect of genetic relatedness, habitat of provenance, and mother tree status on seedling traits was analysed. Nine traits displayed significant genetic effects, while mother tree status and habitat effects were not significant (P > 0.05) for an y trait. Estimated heritability varied between 0.14 and 0.28, with growth-related traits having the highest values. Additive genetic variation correlated positively with nonheritable variation, suggesting that ecological–evolutionary factors increasing or decreasing additive genetic variance may also affect nonheritable variation in the same direction. Our results suggest that quantitative genetic variability should be taken into account in ecological studies on, and in the management of, natural tropical rainforests; further research is needed to investigate genetic × environment interactions, in particular from the point of view of the genetic response of shade-tolerant plant species to variations in light availability.     

Basic density of radiata pine in New Zealand: genetic and environmental factors

Wood basic density is among the selection criteria for many fast-grown tree species, including Pinus radiata D. Don in New Zealand. Basic density was assessed in 23,330 stem cores from 18 trials to study the heritability, the relevance of environmental effects and the magnitude of genotype-by-environment (GxE) interaction. Site differences in annual average temperature dominated variability in this dataset, with lower latitude and altitude (i.e. warmer) sites displaying higher average density. Between highest- and lowest-density sites there was an 18% difference (302.7 vs. 358.4 kg m^− 3) for the linear mean for cores of rings 1–5 and a 39% difference (329.7 vs. 459.1 kg m^− 3) for the linear mean of rings 6–10. The estimated heritabilities fluctuated between 0.28 and 0.94 (mean, 0.6); however, basic density displayed little within-site variability (phenotypic coefficient of variation, <8%). Bivariate analyses were used to estimate between-site genetic correlations as an indication of GxE interaction. Only 57 out of the 153 pairs of trials contained enough information to estimate the between-site genetic correlations and, out of those, 15 estimates were not statistically significant. Moderate to high (0.46–0.96) significant genetic correlation estimates indicated that there was little interaction for basic density, suggesting no need to modify the breeding strategy to account for differential performance in this trait. Poor connectedness between trials could be depressing estimates of genetic correlations. This situation should be considered when designing genetic testing schemes, particularly when purposely inducing imbalance as in rolling front strategies.     

High heritability for heartwood in north Swedish Scots pine

 In 44-year-old full-sibs of north Swedish Scots pine (Pinus sylvestris L.), the estimated heritability of heartwood diameter was 0.5 despite the influence of environmental changes (caused by an earlier thinning) which apparently had reduced the heritabilities of height and diameter to around zero. The heritability of bole straightness was estimated to be 0.6. The coefficient of additive genetic variation of heartwood diameter was estimated at 0.2. If a reliable ‘heartwood-meter’ becomes available that allows nondestructive measurements to be rapidly made in the field it should be possible to breed for or against heartwood formation with less effort compared with that required in breeding aimed at improving regular production traits. Received: 10 June 1998 / Accepted: 8 September 1998     

Genetic variation in basic density and modulus of elasticity of coastal Douglas-fir

Douglas-fir trees from 39 open-pollinated families at four test locations were assessed to estimate heritability of modulus of elasticity (MOE) and basic density. After trees were felled, sound velocity was measured on 4-m logs with the Director HM200. Disks were taken to estimate dry and green wood density; dynamic MOE was estimated as green density × (sound velocity)². Heritability estimates of MOE (across-site h ²=0.55) were larger than those for total height (0.15) and diameter at breast height (DBH; 0.29), and similar to those for density (0.59). Negative genetic correlations were found for MOE with height (r _A=−0.30) and DBH (r _A=−0.51), and were similar to those found for density with height (r _A=−0.52) and DBH (r _A=−0.57). The partial correlations of height with MOE and density, while holding DBH constant, were positive, implying that the observed negative correlations between height and the wood properties were a function of the high positive correlation between height and DBH and the strong negative correlations between DBH and the wood properties. Taper [DBH/(height−1.4)] was found to be negatively associated with MOE. Selection for MOE may produce greater gains than selection for density because MOE had a larger coefficient of additive variation (9.6%) than density (5.1%). Conversely, selection for growth may have a more negative impact on MOE than density because of the greater genetic variation associated with MOE. Family mean correlations of the wood quality traits with stem form and crown health were mostly nonsignificant.     

Provenance variation and genetic parameters of Eucalyptus viminalis in Argentina

Genetic parameters for growth, stem straightness, pilodyn penetration, relative bark thickness and survival were estimated in a base-population of five open-pollinated provenance/progeny trials of Eucalyptus viminalis. The trials, located in northern, central and southern Buenos Aires Province, Argentina, comprised 148 open-pollinated families from 13 Australian native provenances and eight local Argentinean seedlots. The Australian native provenances come from a limited range of the natural distribution. Overall survival, based on the latest assessment of each trial, was 62.4%. Single-site analyses showed that statistically significant provenances differences (p?<?0.05) for at least one of the studied traits in three out of the five trials analyzed. The local land race performed inconsistently in this study. The average narrow-sense individual-tree heritability estimate $ \left( {{{\hat{h}}^2}} \right) $ was 0.27 for diameter and 0.17 for total height. Values of $ {\hat{h}^2} $ also increased with age. Pilodyn penetration, assessed at only one site, was more heritable $ \left( {{{\hat{h}}^2} = 0.32} \right) $ than the average of growth traits. Estimated individual-tree heritabilities were moderate to low for stem straightness (average of 0.20) and relative bark thickness (0.16). The estimated additive genetic correlations $ \left( {{{{r}}_{{A}}}} \right) $ between diameter and height were consistently high and positive ( $ {{r}_{^A}} $ average of 0.90). High additive genetic correlations were observed between growth variables and pilodyn penetration ( $ {{r}_{^A}} $ average of 0.58). Relative bark thickness showed a negative correlation with diameter $ \left( {{{{r}}_{^A}} = - 0.39} \right) $ and height $ \left( {{{{r}}_{^A}} = - 0.51} \right) $ . The average estimated additive genetic correlation between sites was high for diameter (0.67). The implications of all these parameter estimates for genetic improvement of E. viminalis in Argentina are discussed.     

Genetic variation of growth and tree quality traits among 42 diverse genetic origins of Tectona grandis planted under humid tropical conditions in Sabah, East Malaysia

Forty-two different genetic origins of teak (Tectona grandis) comprising 26 open-pollinated families from a clonal seed orchard (CSO) were planted in a replicated trial under 2,500 mm of annual rainfall and no distinct dry season, in 1997, in Sabah, East Malaysia. The trees were measured or scored for various traits at 13, 35, 49, 61, 72, 85, 96, and 106 months after planting. Mortality rate, height (H), diameter at breast height (DBH), volume (V), and fork height (FH) varied strongly among populations and origins. The best population means after 106 months for growth H (21.1 m), DBH (21.1 cm), and V (278 dm³) were for the CSO families. Narrow sense heritabilities for the CSO families increased gradually with age but remained lower after 106 months for DBH (h ² = 0.24) and V (h ² = 0.34) than for H (h ² = 0.51) and FH (h ² = 0.56). Overall, the CSO families were also straighter, less forked, and grew more vertically than the native provenance and seed-derived sources. Such differences did not exist for flowering ability, and at 106 months, the great majority of the trees of the various origins had not yet entered the flowering stage. Overall, at 106 months, the phenotypic correlations between the various quantitative and qualitative traits were weak, except between straightness and bending with values higher than 0.50. These findings confirm the usefulness of CSO for teak improvement and demonstrate the beneficial influence of wet tropical conditions on traits of major economical importance for this species.     

Genetic structure of an introduced paper wasp,Polistes chinensis antennalis (Hymenoptera,Vespidae) in New Zealand

Several eusocial wasps are prominent invaders to remote islands. The paper wasp Polistes chinensis antennalis is native to East Asia, was introduced to New Zealand in 1979 and has expanded its distribution there. This provides an excellent opportunity to examine the impacts of an initial bottleneck and subsequent expansion on genetic structure. We analysed and compared the genetic population structures of the native (Japan and South Korea) and invasive New Zealand populations. Although 94% of individuals had shared haplotypes detected across both populations, the remaining 6% had private haplotypes identified in only one of the three countries. The genetic variation at microsatellite loci was lower in New Zealand than in native countries, and the genetic structure in New Zealand was clearly distinct from that in its native range. Higher frequencies of diploid‐male‐ and triploid‐female‐producing colonies were detected in New Zealand than in the native countries, showing the reduction in genetic variation via a genetic bottleneck. At least two independent introductions were suggested, and the putative source regions for New Zealand were assigned as Kanto (central island) and Kyushu (south island) in Japan. Serial founder events following the initial introduction were also indicated. The estimated dispersal distance between mother and daughter in New Zealand was twice that in Japan. Thus, the introduction history of P. chinensis antennalis in New Zealand is probably the result of at least two independent introductions, passing through a bottleneck during introduction, followed by population expansion from the point of introduction.     

Paternity analysis in a progeny test of Cryptomeria japonica revealed adverse effects of pollen contamination from outside seed orchards on morphological traits

To evaluate the effects of pollen contamination from outside of Cryptomeria japonica seed orchard on the growth performance (height and diameter at breast height, DBH) and morphological traits (stem straightness and basal stem straightness), paternity testing using seven microsatellite markers was performed in a progeny test. In the studied progeny test, high rates of inconsistency were found between the observed and expected genotypes. The average rates of pollen contamination from outside the orchard and self-fertilization were 58.47% and 0.65%, respectively. We divided the individuals of the studied progeny test into two groups based on their genotype data, for which: (1) both parents were elite trees and (2) only the mother trees were elite trees, and then compared them with respect to the growth performance and morphological traits of progenies using data at 20 and 30 years old. Significant adverse effects of contaminating pollen were detected in relation to straightness, but not tree height and DBH. The results suggest that the genetic gains for straightness generally show higher narrow-sense heritability than growth traits, which should be increased by reduction of pollen contamination. Breeding with paternal analysis (BWPA) is an effective approach for evaluating breeding materials based on maternal and paternal information revealed by DNA markers. The use of BWPA in progeny test allows effective forward- and backward selection without laborious and time-consuming tasks. In this study, we also suggest that the significant pollen contamination and paternal deviation found in the open-pollinated progeny test are serious impediments for BWPA.     

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.     

Molecular phylogenetics of the genus <Emphasis Type="Italic">Physalia</Emphasis> (Cnidaria: Siphonophora) in New Zealand coastal waters reveals cryptic diversity

Physalia is a genus of pelagic colonial hydrozoans often known by common names such as ‘Portuguese-man-of-war’ or ‘bluebottle’. Siphonophore systematists generally recognise only a single species in this genus, Physalia physalis, however the name Physalia utriculus is also still in common use, which has led to considerable taxonomic confusion. With some morphological variation between global regions there is the possibility that this genus holds a substantial amount of cryptic variation. We seek to examine the genetic structure of Physalia present in New Zealand coastal waters. Fifty-four specimens collected from 13 locations around New Zealand and Australia were sequenced for both mitochondrial cytochrome c oxidase I (COI) and the first internal transcribed spacer (ITS1) of the nuclear ribosomal cistron. Sequences were analysed using maximum likelihood and split decomposition neighbour networks to determine conflict between clans (the unrooted analog of clades). Three clans were identified from both the COI and ITS sequences. The results are complex and clans are not consistent between the two genes. Nevertheless, it seems that there is substantial cryptic diversity amongst Physalia present in New Zealand coastal waters.     

Few genetic differences between Victorian and Western Australian blue penguins,Eudyptula minor

Abstract

Blue penguins, Eudyptula minor, breeding on Penguin Island, Western Australia are considerably larger than other blue penguins in Australia. If genetic isolation is the cause, it may have implications for the conservation status of some blue penguin populations. We compared the sequences of two mitochondrial gene regions (cytochrome‐b and the control region) from Western Australian blue penguins with other populations of blue penguins from Australia and New Zealand. We found few differences between sequences from Western Australia, Phillip Island, Victoria and Otago, New Zealand, although all three differed considerably from other New Zealand blue penguins. Sequences for the control region from the Western Australian blue penguins and 30 more birds breeding at various Australasian sites provided further support for two major clades within Eudyptula; an Australian clade (including Otago) and a New Zealand clade.