Genetic parameters and genotype by environment interaction of Eucalyptus grandis populations used in intraspecific hybrid production in South Africa

In South Africa, Eucalyptus grandis is an important species due to its fast growth and general suitability of its timber for a range of products. However, E. grandis is susceptible to fungal diseases such as Crysoporthe austroafricana and Coniothyrium sp. cankers in the subtropical region of Zululand and is therefore mainly planted as a parental species in a hybrid combination with E. urophylla in this region. The current strategy is to maintain large breeding populations of both parental species in order to provide improved elite selections for hybrid crosses. In order to develop the best interspecific hybrid breeding strategy for E. grandis, it is important to first determine estimates of genetic parameters of the pure species parents. Estimating the genotype by environment interaction (G×E) is also necessary in proposing the basis for setting up breeding populations and selecting environmentally stable genotypes. With this in mind, two E. grandis full-sib progeny trials were planted in Zululand and one in the KwaZulu-Natal Midlands region. The aims of this study were firstly to determine the magnitude of G×E of E. grandis across the three sites; secondly, to estimate the genetic parameters for growth of the E. grandis parents selected for intraspecific crosses; and lastly, to identify the best parents to use for intra- and interspecific crosses in future hybrid breeding programmes. Results of our study indicated that G×E would be practically negligible for growth in Zululand and one group of elite parents can be used for hybrid crosses in this region. In general, growth traits were under low to moderate genetic control, and the variation in additive genetics enabled us to identify E. grandis parents that could be utilised for intraspecific crosses and deliver progeny with genetic gains of 28.4%. Our study also highlighted that a relatively large portion of the genetic variation was explained by dominance genetic variation and a strategy to capture this non-additive variation needs investigation. Although our study achieved the stated aims, it must be kept in mind that E. grandis is mainly used as a hybrid parent with E. urophylla in Zululand. A study to investigate whether the parents with good general combining ability values from our study are also good general combiners in interspecific hybrid combinations with E. urophylla needs to be conducted.     

A comparison of deterministically predicted genetic gains with those realised in a South African Eucalyptus grandis breeding program

Tree breeders attempt to predict the genetic gains that are likely to be achieved through selection and breeding of new generations, using stochastic or deterministic modelling. There are many factors that may cause a discrepancy between the predicted and realised genetic gains. Often the predictions for genetic gains are based on single trait selection, whereas in reality the breeding tends to be multitrait in nature. The violation of Hardy-Weinberg conditions, assumptions regarding outcrossing and relatedness, assumptions regarding the effect of the interaction between the environment and the genotype, and numerous possible errors in the process of breeding, all could result in unexpected discrepancies between the realised and predicted genetic gains. A series of genetic gains trials containing representatives of three generations of Eucalyptus grandis selections were compared with the view to verifying the effectiveness of the E. grandis breeding program. Genetic gains of the F₃ (third generation of pedigreed progeny) over the F₂ generation (second generation of pedigreed progeny) were 15% for tree growth (volume). A comparison between F₂ and P₀ revealed an improvement of between 20% and 33% for growth. This exercise highlighted complexities of modelling the predicted genetic gains of assimilated genetic breeding trials. The predictions of genetic gains did deviate (in both directions) from those realised, although these deviations may be explained as functions of imperfect modelling. On average, however, the predicted genetic gains for tree volume over three generations was 13% between generations, whereas the average realised genetic gain in the genetic gains trial was 14%. It is therefore assumed that the E. grandis breeding population is indeed performing as expected, following classical tree breeding assumptions.     

Genetic control of wood density and bark thickness,and their correlations with diameter,in pure and hybrid populations of Eucalyptus grandis and E. urophylla in South Africa

Tree diameter under and over bark at breast height (dbh), wood density and bark thickness were assessed on samples from control-pollinated families of Eucalyptus grandis, E. urophylla, E. grandis × E. urophylla and E. urophylla × E. grandis. The material was planted in field trials in the coastal Zululand region of South Africa. At 75 months, between three and seven of the best trees per family were felled and wood samples collected. Genetic parameters for wood density, bark thickness and bark percentage (ratio of double bark thickness to overbark diameter) and the inter-trait correlations for the different species and hybrids were calculated. Genetic parameter estimates for wood density, bark thickness and bark percentage in the E. urophylla × E. grandis hybrids showed these traits to be under total additive genetic control. This was confirmed by the intermediate hybrid means for these traits relative to those of the parental species. There was a very low correlation between dbh and wood density for the E. urophylla × E. grandis hybrids (r_G = –0.07 and r_P = 0.064). Amongst the E. urophylla families there was a moderate positive and significant phenotypic correlation between wood density and bark thickness (r_P = 0.391), and between wood density and bark percentage (r_P = 0.442).     

Genetic variability of growth and wood chemical properties in a clonal population of Eucalyptus urophylla × Eucalyptus grandis in the Congo

Thirty-four Eucalyptus urophylla × Eucalyptus grandis hybrids were evaluated with a view to selecting for improved growth and wood-quality traits for plantations in the Congo. Height, circumference at breast height and volume were measured at 12, 27, 37, 49 and 60 months. Lignin content, the syringyl/guaiacyl ratio and total extractives content were predicted by near-infrared spectroscopy using wood powder samples collected from trees at breast height. While wood chemical properties were stable and under strong genetic control, growth traits were not. The genetic correlation between lignin content and growth was weak and negative, whereas the environmental correlation was also weak but positive. The genetic improvement of E. urophylla × E. grandis clones, based on growth features, leads to a limited decrease in lignin content and syringyl content and to a limited increase in extractives content.     

Realised genetic gains and estimated genetic parameters of two Eucalyptus grandis × E. urophylla hybrid breeding strategies

Conventionally, Eucalyptus grandis × E. urophylla (GU) hybrid material is first tested as seedlings in progeny trials for at least four years before ortets are selected and ramets of the selected ortets are propagated to test in clonal trials. The primary constraint with this ‘conventional hybrid breeding strategy’ (CHBS) is the time required to first test the hybrid material as seedlings. In order to address this, an ‘accelerated hybrid breeding strategy’ (AHBS) was investigated to reduce the time spent on testing GU hybrid material as seedlings. However, it is of utmost importance to quantify the impact the AHBS might have on genetic gains and genetic information. With this in mind, two clonal populations have been established with genetic material that derived from the CHBS and the AHBS. The main purpose of this study was to do a comparative study between the CHBS and AHBS, firstly to quantify the genetic gains per unit time for GU hybrid clonal populations that have been derived from the CHBS and AHBS respectively, and secondly to obtain genetic parameters such as heritabilities, the ratio of dominance, clonal within family variance, and the proportion of additive and non-additive genetic variance. The results indicated that the percentage realised volume gains per year was higher for the AHBS (3.7%) than for the CHBS (1.9%) when compared with the GU commercial clone. Thus, shortening the testing time of GU seedlings had a positive impact on volume gains per year. With regard to genetic parameters, both the AHBS and CHBS clonal populations indicated that non-additive genetic variation explained the majority (88% and 71%, respectively) of the genetic variation. Due to the pre-eminence of non-additive genetic variation, the narrow-sense heritabilties for the female and male effects were negligible for both clonal populations. Overall, the majority of the non-additive genetic variation was explained by the proportion of dominance variance, and less by the clone within family effect. These results suggest that, firstly, the time spent on testing GU hybrid material as seedlings should be minimised and, secondly, a hybrid breeding strategy to capture non-additive genetic variation should be adopted.     

尾叶桉×细叶桉木材密度与生长的联合选择

[目的]评估Resistograph钻刺法间接测定尾叶桉×细叶桉木材密度的可靠性,检测杂交亲本对子代表型的效应以及生长与木材密度的相关,评选速生、优质的尾细桉杂种。[方法]基于10株尾叶桉与10株细叶桉不完全析因交配产生的56个杂交组合的7.5年生试验林,利用79株分析容积法与Resistograph钻刺法测定的木材密度的相关,通过方差分析检测亲本对杂种生长和木材密度的效应,结合多重比较和独立淘汰法进行材积和木材密度的联合选择。[结果]容积法与Resistograph钻刺法测定的木材密度的表型相关系数为0.52(P0.001),遗传相关系数为0.55(P0.05);树高、胸径及材积的母本间和父本间均呈极显著差异(P0.001或0.01),但母本×父本互作的效应不显著;对钻刺木材密度,父本间呈极显著差异(P0.001),母本×父本互作显著(P0.05),但母本间差异不显著;树高、胸径和材积间的表型相关和遗传相关均极显著(P0.001),其与钻刺木材密度的表型相关极显著(P0.001),但遗传相关不显著;评选出速生、木材密度较高的杂交组合14个、单株17株。[结论]Resistograph钻刺法是一种间接测定尾细桉木材密度的简便、经济和可靠的方法;母本和父本选择以及母本与父本的组配对培育速生、材质优良的尾细桉杂种均较重要;尾细桉生长与木材密度的遗传相关不显著,需要对这两类性状分别进行选择;评选的尾细桉杂交组合和单株为培育速生、优质的桉树良种提供了有用的材料。     

桉属树种与种源、家系比较试验

以９个桉属树种，其中８个种泊来自原产地，７个家系和７个杂种后代来自国内次生资源进行混合比较，５年生评价结果表明：树种、种源、家系呈极显著差异；尾叶桉及其两个种源、尾巨桉杂交种细叶桉的１３５４４种源适合本试验区戒严和格短轮伐期经营，可获得适应性和生长指标平均值超过平均水平３５％的选择效应。其中最优为１４５３４号尾叶桉种源，５年生单株材积０．０５０．６１ｍ＾３／株（８９．０３ｍ＾３／ｈｍ＾２）。种子采     

Genetic parameter estimates for interspecific Eucalyptus hybrids and implications for hybrid breeding strategy

The Forestry Commission of Zimbabwe initiated a hybrid breeding program of Eucalyptus grandis with E. tereticornis (G × T) and E. camaldulensis (G × C) in order to provide suitable genotypes for planting in areas that are marginal for E. grandis in terms of drought and frost. A total of seven hybrid trials were established at four sites, representative of low to medium rainfall. The female parents were from E. grandis preselected for superior volume production. No pure species were included in the trials to act as controls and allow for estimation of hybrid vigour. Survival, growth and stem straightness were assessed at 18 and 43 months from planting. Survival was generally above 80% for all hybrids across sites. G × C hybrids performed better than G × T hybrids, showing a 20% difference in mean height at 43 months. Female and male variances were generally insignificant (P > 0.05) and in some cases, the estimates were 0. This was not unexpected, as the parents for the hybrids were preselected for superior height, diameter at breast height (DBH) and volume production as pure species. Ratio of dominance to phenotypic variance ( $ \hat{d}^{2} $ ) was significant for all traits for all hybrids (P < 0.05). Trait–trait dominance correlations (r _D ) at 43 months between height and DBH were generally large and positive, above 0.6. Type-B dominance correlation (r _BD ) suggested evidence of genotype-by-environment interaction (G × E), but one high elevation site contributed most to the observed G × E. If these results are confirmed in larger mating designs with progeny established on multiple sites, it seems that the most appropriate breeding strategy for E. grandis × E. tereticornis and E. grandis × E. camaldulensis hybrids may be one that exploits both additive and dominance genetic variance, such as the reciprocal recurrent selection (RRS) scheme or RRS with forward selection.     

桉属树种/种源试验

广东省江门地区为调整人工林树种结构 ,提高人工林的产量和质量 ,发展以阔叶树种为主的人工林。现桉树 ( Eucalyptusspp.)已成为该市营造短轮伐期工业用材林的主要树种 [1] 。本试验的目的在于通过从澳大利亚引进桉属树种 ,进行桉属树种 /种源试验 ,为江门市选出更适合当地生长的优良桉树树种和优良种源 ,促进该地区桉树人工林的发展。1　试验地概况　　试验地位于广东省开平市镇海林场 ,2 2°2 5′～ 2 2°40′N,112°15′～ 112°45′E。属南亚热带气候 ,年平均温度 2 2 .1℃ ,极端最高温度 38.3℃ ,极端最低温度 1℃ ,年平均降水量 182…     

Quantitative genetics of cold hardiness and growth in <Emphasis Type="Italic">Eucalyptus</Emphasis> as estimated from <Emphasis Type="Italic">E.</Emphasis><Emphasis Type="Italic">urophylla</Emphasis> × <Emphasis Type="Italic">E. tereticornis</Emphasis> hybrids

Interspecific hybrids of Eucalyptus urophylla S. T. Blake × E. tereticornis Smith were used for quantitative genetic analysis of cold hardiness (CH) and growth traits height (H), diameter at breast height (D) and volume (V) at 0.5, 1.5, 2.5 and 4 years of age. The effect of E. tereticornis male variance was significant (P < 0.05) for CH as well as 2.5- and 4-year-old growth except for H ₄, and E. urophylla female × E. tereticornis male interaction was significant for 0.5- and 1.5-year-old growth and CH except for CH _0.5, whereas E. urophylla female effect was not significant for the majority of traits analyzed. Both additive and dominance gene action were present in the genetics of all the traits measured at most ages, indicating that those traits may be improved by hybrid breeding. The σ _D ²/σ _A ² ratios tended to decline after 1.5 years of age in CH and growth traits, implying that breeding value predictions derived from older performance might be more reliable. Estimates of narrow-sense heritability (h ² ) for CH and growth ranged from 0.03 ± 0.05 to 0.28 ± 0.13, depending on trait and age. Additive genetic correlations between CH and V were 0.28 ± 0.48 and 0.69 ± 0.30 at ages 1.5 and 2.5, respectively, suggesting that selection for growth would cause a favorable increase in cold hardiness in E. urophylla × E. tereticornis hybrids.     

桉树蛀干新害虫--桉蝙蛾形态与生物学研究

桉蝙蛾是近年来在广西、广东新发现严重危害桉树的蛀干害虫,本文记述了该害虫的形态特征和生物学特性,该虫在广西分布于12个市49个县,10个自治区直属国有林场,232个乡镇,危害桉树面积达2 894.42 hm2,多数个体1年完成1代,以幼虫于12月中下旬开始在受害树干中越冬,次年2-3月化蛹,4月羽化,5月上旬初孵幼虫出现,6-8月幼虫上树转移危害,1年只有1次羽化;该虫主要分布在丘陵山地,以沟谷和坡底危害较严重,主要危害1～2年生巨尾桉和尾巨桉无性系.1年生和2年生受害桉树单株材积生长量下降率分别为16.70％和24.78％.沟谷和坡底1～2年生速生桉无性系是防治的主要对象.     

Ralstonia solanacearum decreases volumetric growth of trees and yield of kraft cellulose of Eucalyptus spp.

Ralstonia solanacearum is currently one of the most important plant pathogenic bacteria worldwide, with a wide geographical distribution and host diversity. The pathogen infects more than 200 plant species belonging to approximately 50 plant families, including Eucalyptus spp. Although, high losses have been reported in nurseries, little is known on the negative impact of the disease in the field. In this study, we evaluated the incidence of R. solanacearum and its effect on volumetric growth and cellulose yield of discoloured wood chips obtained from infected trees of one clone of Eucalyptus urophylla and two hybrid clones of Eucalyptus urophylla × E. grandis. The average incidence of bacterial wilt ranged between 60.6% and 72.4%. Volumetric growth of infected trees disease decreased 78.6% and 81.7% at 18 and 30 months, respectively. The pulp screen yield of three clones decreased between 3.2 and 6.4%, with an average 4.3%. The results of this work provide useful information on the losses of volumetric growth and pulp yield of eucalypt caused by R. solanacearum.     

The Eucalyptus shoot and leaf pathogen Teratosphaeria destructans recorded in South Africa

Species of Teratosphaeria include some of the most important fungal pathogens of plantation-grown eucalypt trees. During routine disease surveys, symptoms and signs of leaf spot and blight were observed on the foliage of one-year-old E. grandis × E. urophylla hybrids in the Zululand region of KwaZulu-Natal, South Africa. These were distinct from those caused by the well-known and leaf-infecting fungus Teratosphaeria suttonii, which is not considered an important pathogen in the country. Culture and morphological characteristics as well as DNA sequences for three gene regions were used to compare the fungus isolated from the newly emerging symptoms with those for known Teratosphaeria species. DNA sequences were the same as those for T. destructans and this was consistent with the distinctive morphology of the asexual spores and the symptoms on leaves. Teratosphaeria destructans is an aggressive pathogen and actions will be needed to ensure that it does not impart serious losses to the local forestry industry.     

不同种植年限尾巨桉人工林叶片-凋落物-土壤碳氮磷化学计量特征

[目的]为了解雷州半岛尾巨桉速生人工林生态系统的C、N、P分配格局及化学计量特征。[方法]采用空间换时间的方法,选取雷州半岛4种不同林龄(1、3、5、7 a)的尾巨桉人工林为研究对象,对尾巨桉叶片、凋落物及土壤的C、N、P含量及化学计量特征进行测定分析。[结果]表明:C、N含量表现为叶片凋落物土壤,P含量表现为叶片土壤凋落物,且3个库间差异显著;土壤的C含量随林龄增加而增加,N、P含量差异不显著,土壤C∶N随林龄的增加而增加,说明土壤有机质分解速率逐渐下降;凋落物的C∶N为54. 07 92. 18 ( 25),表明尾巨桉林下凋落物分解速率较慢,N元素成为主要限制凋落物分解的元素,凋落物的C∶N随林龄的增加先增加后下降,凋落物分解速度先降低后升高;叶片的N∶P为10. 80 12. 98,说明中幼林龄尾巨桉受N限制较明显。相关性分析表明:凋落物养分元素含量受叶片限制,土壤养分含量受凋落物限制,表明生态系统内部C、N、P元素在植物、凋落物与土壤之间实现了运输和转换。[结论]雷州半岛尾巨桉中、幼林龄时期土壤有机质及凋落物分解速率较慢;随林龄的增加,土壤有机质、凋落物分解速率下降,N元素成为其主要分解限制性元素,林分生长受N限制明显。     

Stomatal characteristics of Eucalyptus grandis clonal hybrids in response to water stress

This study describes the stomatal response occurring during water stress and subsequent recovery of three Eucalyptus grandis clonal hybrids. The aim was to investigate the degree to which stomatal conductance (g _s) and stomatal density differ between the clonal hybrids across seasons and in response to water stress. Plants from one E. grandis × E. camaldulensis (GC) and two E. grandis × E. urophylla (GU1 and GU2) clones were grown for 18 months in 80 l planting bags. Plants were subjected to three watering treatments: control (100% field capacity), chronic water stress (maintained at 15% of field capacity) and acute water stress (cyclic water stress, where water was withheld until leaf wilting point, and a subsequent period of recovery followed). Stomatal conductance was measured after 6, 12 and 18 months growth. At 12 months of age, the recovery of g _s 1, 2 and 7 d after rewatering (following acute water stress) was further investigated. The GC hybrid showed consistently higher g _s than the GU clones at each measurement period. Stomatal conductance was 24–66% higher during winter (after 12 months growth) than during summer. The recovery of stomatal conductance from acute water stress was more rapid in the GC clone than the GU clones. Chronic water stress was shown to decrease g _s in GU clones by up to 70%, but not in the GC clone. Water stress did not affect stomatal density or size. Remarkably, stomata were absent from the adaxial leaf surface of clone GU1 leaves, but not from the leaves of the other E. urophylla hybrid cross (GU2). Total biomass of the GC clone was significantly greater at 9 months growth, but after 18 months growth the GU1 clone had attained greater biomass accumulation (although not significantly). Measurement of g _s, transpiration, stomatal density and total biomass in the GU1 clone indicated stomatal sensitivity to water stress, a favourable trait during periods of drought. The differing growth strategies of the GU and GC clones could be partially explained by their differences in stomatal sensitivity in response to water stress.     

Dissimilar stem and leaf hydraulic traits suggest varying drought tolerance among co-occurring Eucalyptus grandis × E. urophylla clones

In South Africa, there is currently a phenomenon where a Eucalyptus grandis × E. urophylla clone is dying on a large scale, whereas co-occurring clones in the same plantation area remain healthy. No primary biotic pathogen has been isolated from these dying trees. However, the plantation region has endured severe drought conditions. We hypothesised that the specific E. grandis × E. urophylla clone dieback is due in part to the affected trees having an elevated drought vulnerability. We tested this hypothesis retrospectively by examining how the dying (branch dieback started) as compared with two healthy and co-occurring E. grandis × E. urophylla clones vary in branch xylem anatomy (water transport) and leaf stable carbon isotopes (gas exchange) across two levels of mean annual precipitation in the planted landscape. There was significant intra-hybrid variation in the range of studied hydraulic traits. It appeared that the dying clone is maladapted to drier field conditions relative to the two healthy growing clones. Individuals of the drought-susceptible clone had increased hydraulic conductivity, having both wider vessel lumens and larger lumen fractions, and also did not regulate leaf-level transpiration in the drier landscape as efficiently as the two unaffected clones (lower δ¹³C ratios). In turn, one of the healthy growing clones had a completely different hydraulic strategy, having higher wood density, lower lumen fractions, and higher δ¹³C ratios that are widely considered as hydraulically resilient during water stress. The other clone appeared to also gain drought tolerance via higher leaf-level water-use efficiency, although drought tolerance was less clear regarding xylem anatomy. We highlight how variation in relative drought vulnerability can be highly significant even between clones within a single hybrid group. Knowledge of this variation in plant hydraulics between commercially planted and closely related Eucalyptus trees would complement clone deployment programmes in a target landscape.     

广西丘陵台地巨尾桉、尾叶桉(实生)地位指数表的编制*

从广西南部桉树主要栽培区丘陵台地立地类型现有林人中收集的９５株解析木及９１株固定标准地内平均优势木调查材料，选用ＬｎＹ＝Ａ＋Ｂ×Ｘ＾ｃ公式模拟出巨尾桉和尾叶桉的导向曲线。最后确定了广西丘陵台地两种桉树的立地指数表。经检验，达到了较高的估测精度。     

Variation in growth potential between hybrid clones of <Emphasis Type="Italic">Eucalyptus</Emphasis> trees in eastern South Africa

Growth of commercial forestry is highly dependent on the availability of fast-growing planting materials. Consequently, the efficient utilization of fast-growing plantations can greatly impact productivity. The objectives of this study were to evaluate variations in the growth potential of two clones and to estimate the average stem radial growth advantage of a fast-growing clone using data obtained from Sappi landholdings in eastern South Africa and a mixed modelling approach that permits the incorporation of covariance structure into the statistical model. During the first 2 years of growth, the stem radius of nine trees each of two clones was measured using dendrometer attached to the tree. A second-degree fractional polynomial model was chosen to show the functional relationship between stem radius and tree age. Growth of the two hybrid clones differed significantly. The Eucalyptus grandis × Eucalyptus urophylla clone grew faster than the E. grandis × camaldulensis clone, indicating better genetic potential for rapid growth and yield. This study can be considered as starting point to further compare the potential for rapid growth of several hybrid clones using the longitudinal data modelling approach.     

速生乡土树种黎蒴优树半同胞家系子代测定

将采自广东的278个黎蒴优树家系种植在广东东江林场和梅南林场,进行半同胞子代测定。造林5年后,对参试家系的3个主要生长性状进行调查分析。黎蒴家系树高、胸径和材积存在显著差异,遗传变异系数在13．60％～67．13％,且变异主要受遗传控制,其家系遗传力分别为0．511～0．543,0．585～0．736和0．469～0．717;相关性分析发现,材积生长量主要受胸径影响,其相关系数大于0．80。在10％～30％入选条件下,各生长性状均具有较高的遗传增益,其中材积遗传增益〉胸径遗传增益〉树高遗传增益。采用育种值评估法与综合指数法共选出优良家系36个、优良单株71株。研究结果为进一步开展黎蒴选育和遗传分析奠定了坚买基础。     

黧蒴栲半同胞家系生长性状变异规律及遗传参数估算

为探讨黧蒴栲半同胞家系生长性状遗传变异规律和估测其遗传参数,采用完全随机区组设计,对52个6年生黧蒴栲半同胞家系进行生长测定分析。结果表明：黧蒴栲家系间胸径、树高和材积生长均存在极显著差异。52个黧蒴栲半同胞家系平均胸径为8.53em,变异幅度为6.31～10.85am;平均树高为8.23m,变异幅度为5.08～10.14m;平均材积为0.02828 m^3,变异幅度为0.01099～0.05407 m^3。黧蒴栲半同胞家系胸径遗传变异系数为13.74％、广义遗传力为0.7404;树高遗传变异系数为13.74％,广义遗传力为0.7150;材积遗传变异系数为35.36％,广义遗传力为0.7539。表明黧蒴栲半同胞家系胸径、树高变异较小,受较强遗传控制,材积变异较大,也受较强遗传控制,通过一定强度的选择,能获得较高的遗传增益。