赤桉抗風(fēng)特性及其重要性狀的關(guān)聯(lián)分析
發(fā)布時(shí)間：2019-07-09 05:58
【摘要】：我國南方沿海夏季臺(tái)風(fēng)頻繁,風(fēng)害嚴(yán)重影響沿海地區(qū)的生態(tài)安全以及相關(guān)產(chǎn)業(yè)的發(fā)展,尤其對林木生長構(gòu)成極大威脅,成為制約桉樹人工林可持續(xù)發(fā)展的重要因素之一。選育優(yōu)良抗風(fēng)品系是保證我國沿海地區(qū)桉樹商品林可持續(xù)發(fā)展的重要措施。赤桉(Eucalyptus camaldulensis)為優(yōu)質(zhì)的硬木材,耐旱、耐瘠、抗風(fēng)、抗病能力強(qiáng),為我國桉樹樹種改良的重要遺傳材料。本論文以114個(gè)赤桉家系(半同胞)為材料對其生長、材性、物理力學(xué)、化學(xué)成分等26個(gè)性狀進(jìn)行跟蹤調(diào)查,對各個(gè)性狀進(jìn)行遺傳變異分析并通過相關(guān)性分析尋找影響桉樹抗風(fēng)的主要影響因素。通過風(fēng)害模擬(拉樹實(shí)驗(yàn))分析風(fēng)與重載荷下的變形以及抗風(fēng)主要影響因子,并對赤桉家系抗風(fēng)特性進(jìn)行評(píng)估,最終獲得影響赤桉抗風(fēng)的主要影響因子及選育出優(yōu)良抗風(fēng)家系。桉樹抗風(fēng)性是由眾多復(fù)雜因素引起的,本研究在獲得影響桉樹抗風(fēng)的主要影響因素的基礎(chǔ)上利用SSR分子標(biāo)記技術(shù)進(jìn)行關(guān)聯(lián)分析,對桉樹抗風(fēng)相關(guān)性狀進(jìn)行關(guān)聯(lián)分析,系統(tǒng)剖析赤桉抗風(fēng)的遺傳基礎(chǔ),發(fā)掘優(yōu)良SSR位點(diǎn)。本研究將有助于建立桉樹抗風(fēng)育種分子標(biāo)記輔助選擇體系,為發(fā)掘和利用重要的桉樹抗風(fēng)基因資源奠定基礎(chǔ),為后續(xù)相關(guān)研究提供理論依據(jù)。主要研究結(jié)論如下:1、赤桉家系各生長性狀間存在著豐富的遺傳變異,家系間性狀各性狀中家系遺傳力均高于單株遺傳力,說明家系的選擇潛力更大一些。早期赤桉(1年生)各家系性狀間存在著豐富的遺傳變異,具有很強(qiáng)的家系和家系內(nèi)單株選擇潛力。風(fēng)害指數(shù)與家系生長性狀的遺傳相關(guān)系數(shù)以冠幅與體積最大,抗風(fēng)性與林木大小相關(guān)性不強(qiáng),回歸方程也表明抗風(fēng)性與其生長性狀關(guān)聯(lián)程度不大,相反林木植株越速生(即材積越大),更易遭受風(fēng)害。2~3生赤桉抗風(fēng)性研究表明,平均風(fēng)害指數(shù)為2級(jí)以下,林木的風(fēng)害指數(shù)與其林木大小(樹高、單株材積)呈強(qiáng)負(fù)相關(guān),不同家系間抗風(fēng)性存在著差異且具有遺傳性,而且某些性狀遺傳控制水平相對較高。3~4年生赤桉家系風(fēng)害等級(jí)與樹高、冠幅、材積、樹皮厚度的Pearson相關(guān)性呈極顯著負(fù)相關(guān)關(guān)系。說明樹木越高、材積越大越抗風(fēng)。3~4年生赤桉家系抗風(fēng)性研究表明赤桉抗風(fēng)能力與樹高、胸徑、材積有關(guān)。2、木材密度在赤桉家系間、不同家系部位間以及其家系和部位交互間差異性均不顯著,40月生的赤桉木材密度屬于小級(jí),即屬輕木材。根據(jù)木材物理力學(xué)分級(jí)標(biāo)準(zhǔn)劃分赤桉干縮率屬于III級(jí),容易開裂,樹干底部差異干縮值較中部、梢部差異干縮值大,所以底部更容易開裂和變形。赤桉不同家系間力學(xué)性狀(抗彎強(qiáng)度、抗彎彈性模量、順紋抗剪強(qiáng)度、順紋抗壓強(qiáng)度)差異性極顯著,不同部位間抗彎彈性模量差異極顯著,其余指標(biāo)差異性不顯著。研究發(fā)現(xiàn)林木大小(樹高、胸徑、材積)是影響桉樹抗風(fēng)性能的主要因子,其次是材性因子。單株立木風(fēng)害程度與立木大小密切相關(guān),也與立木的材性如纖維寬度、Pilodyn值、應(yīng)力波值有著較強(qiáng)的相關(guān)性。從木材基本密度和干縮率、物理力學(xué)(抗彎強(qiáng)度、抗彎彈性模量、順紋抗剪強(qiáng)度、順紋抗壓強(qiáng)度等)等力學(xué)性狀與赤桉抗風(fēng)性之間相關(guān)性可以看出二者之間相關(guān)性不顯著,風(fēng)害指數(shù)與木質(zhì)素、綜纖維素、α-纖維素、半纖維素之間相關(guān)性均不顯著(P0.05),從相關(guān)性分析發(fā)現(xiàn)赤桉抗風(fēng)性能與木材物理力學(xué)及化學(xué)成分關(guān)系不大。3、進(jìn)一步通過風(fēng)害模擬(拉樹實(shí)驗(yàn))獲得的拉力三因素(立木所受拉力、樹干變形程度及傾斜角度)與生長、材性、物理力學(xué)、化學(xué)成分等26個(gè)性狀進(jìn)行回歸分析及典型相關(guān)分析發(fā)現(xiàn):影響赤桉樹干拉樹三因素的主要影響因子為樹高、材積、胸徑、樹皮厚度、纖維長度、氣干干縮率、抗拉強(qiáng)度、抗拉彈性模量、綜纖維素含量。其中對赤桉抗風(fēng)影響最大的赤桉的生長性狀即樹高、材積、胸徑,其次是纖維長度、抗拉強(qiáng)度。4、遺傳多樣性分析參試材料為8個(gè)赤桉群體的109株樣品,參試標(biāo)記為覆蓋全基因組的107個(gè)SSR位點(diǎn)。55個(gè)中性的基因組SSRs揭示了赤桉群體多樣性水平較高,共檢測到1116個(gè)等位片段,平均每位點(diǎn)20.3個(gè)等位片段,PIC為0.3508~0.9489,平均0.771,位點(diǎn)均為多態(tài)性較高的位點(diǎn),HE平均值為0.718,說明赤桉發(fā)生高度雜合,Fst平均值為0.066,群體間的遺傳分化不顯著,分子方差分析中群體間方差分量僅占1%,說明遺傳變異主要存在于群體內(nèi)。從赤桉Nei(1972)遺傳距離分析發(fā)現(xiàn)澳大利亞亞群遺傳背景差異較大。赤桉群體具有豐富的遺傳多樣性,可進(jìn)行抗風(fēng)品系選育和遺傳改良的基因庫。5、利用800多對覆蓋全基因組的SSR標(biāo)記通過PCR優(yōu)化以及對8株強(qiáng)抗風(fēng)和8株弱抗風(fēng)赤桉基因池的篩選,得到在兩類基因池中等位頻率差異較大的107個(gè)SSR標(biāo)記用于赤桉抗風(fēng)相關(guān)性狀的SSR關(guān)聯(lián)分析。赤桉家系樹高、胸徑、材積、風(fēng)害指數(shù)等4個(gè)性狀與26個(gè)SSR位點(diǎn)存在顯著連鎖(P0.05),可解釋生長及抗風(fēng)性狀表型變異的9.26%~71.14%。其中樹高相關(guān)位點(diǎn)最多,為13個(gè),其他性狀5~8個(gè)。26個(gè)關(guān)聯(lián)位點(diǎn)中EUCeSSR235位點(diǎn)對風(fēng)害指數(shù)的解釋率最高為71.14%,EUCeSSR570位點(diǎn)對樹高解釋率最低為9.26%,平均解釋率為36.27%。位點(diǎn)EUCeSSR332與樹高、胸徑、材積、風(fēng)害指數(shù)等4個(gè)性狀均顯著相關(guān)聯(lián),該位點(diǎn)對4個(gè)性狀表型變異解釋率相對較高,超過60%。位點(diǎn)EUCeSSR484、EUCeSSR352、EUCeSSR570和EUCeSSR422與樹高和風(fēng)害指數(shù)等2個(gè)性狀均相關(guān)聯(lián),EUCeSSR489和EUCeSSR114位點(diǎn)與樹高和材積等2個(gè)性狀相關(guān)聯(lián)。
文內(nèi)圖片：
圖片說明：09份赤桉種質(zhì)資源群體結(jié)構(gòu)圖
[Abstract]:The frequent occurrence of typhoon and wind damage in the coastal areas of the south of China have seriously affected the ecological security of the coastal areas and the development of the related industries, in particular, it poses a great threat to the growth of the trees, and becomes one of the important factors to restrict the sustainable development of the eucalyptus plantation. The breeding of the good wind-resistant strain is an important measure to ensure the sustainable development of the eucalyptus goods in the coastal area of China. Eucalyptus camaldulensis is an important genetic material for the improvement of eucalyptus tree species in China. In this paper,26 characters, such as growth, material property, physical mechanics, chemical composition and so on, were investigated by using 114 red eucalyptus family (half-siblings), and the main influencing factors that affected the wind resistance of the eucalyptus were found through the analysis of the genetic variation and the correlation analysis. The wind and heavy load deformation and the wind-resistant main influence factors are analyzed by the wind damage simulation (Rtree experiment), and the wind resistance characteristics of the Eucalyptus urophylla family are evaluated, and the main influence factors affecting the wind resistance of the Eucalyptus urophylla are obtained, and the excellent anti-wind family is selected. The wind resistance of Eucalyptus is caused by a number of complex factors. This study is based on the main influencing factors that affect the wind resistance of the eucalyptus, and uses the SSR molecular marker technology to carry out the correlation analysis. The correlation analysis of the wind resistance of the eucalyptus is carried out, and the genetic basis of the wind resistance of the eucalyptus is analyzed. And the excellent SSR locus is excavated. This study will help to establish a molecular marker-assisted selection system for wind-resistant breeding of eucalyptus, laying the foundation for exploring and using the important anti-wind gene resources of the eucalypt, and providing a theoretical basis for the follow-up research. The main results of the study are as follows:1. There are abundant genetic variation among the growth characters of the family of Eucalyptus, and the heritability of the family is higher than the heritability of the single plant, and the selection potential of the family is larger. There are abundant genetic variation among the characters of the early Eucalyptus (1-year-old), which has a strong family and single-plant selection potential in the family. The genetic correlation coefficient between the wind-damage index and the growth character of the family is the largest, the resistance to wind resistance and the size of the tree is not strong, and the regression equation also shows that the resistance to wind resistance is not related to the growth character of the family, and the faster the tree plant grows (i.e. the larger the volume product), The results showed that the average wind-damage index was below grade 2, and the wind-hazard index of the forest was negatively correlated with the tree size (tree height and single-plant volume). The genetic control level of some characters was relatively high. The correlation of the wind-damage grade of the 3-4-year-old Eucalyptus family with the tree height, the crown width, the volume of the material and the Pearson correlation of the bark thickness was negatively correlated. It is indicated that the higher the tree, the greater the volume of the material, and the wind resistance of the 3-4-year-old Eucalyptus urophylla family shows that the wind resistance of the eucalyptus is related to the height of the tree, the diameter of the breast and the volume of the tree. There was no significant difference in the inter-family and family and site interactions among the different families, and the wood density of the 40-month-old Eucalyptus grandis belongs to the small grade, which belongs to the light wood. According to the physical and mechanical classification standard of the wood, the dry shrinkage rate of the eucalyptus is grade III, is easy to crack, the dry shrinkage value of the difference between the bottom of the trunk and the middle part of the trunk is relatively large, and the difference in the dry shrinkage of the tip part is large, so that the bottom is more prone to cracking and deformation. The difference of the mechanical properties (bending strength, flexural modulus, the shear strength and the compressive strength) of the different parts of Eucalyptus grandis is very significant, the difference of the flexural modulus of the different parts is very significant, and the difference of the other indexes is not significant. The study found that the size of trees (height, DBH, volume of wood) was the main factor that affected the wind resistance of the eucalyptus, followed by a material factor. The degree of wind damage of a single plant is closely related to the size of the vertical wood, and also has a strong correlation with the wood properties such as the fiber width, the Pilodyn value and the stress wave value. The correlation between the mechanical properties such as the basic density of the wood and the dry shrinkage, the physical mechanics (bending strength, the flexural modulus of bending, the shear strength of the line, the compressive strength of the smooth grain, etc.) and the anti-wind resistance of the eucalyptus urophylla can be seen to show that the correlation between the two is not significant, and the wind-hazard index is similar to that of the lignin and the heddle, The correlation between cellulose and hemicellulose was not significant (P0.05). From the correlation analysis, it was found that the anti-wind property of Eucalyptus globulus was not related to the physical and chemical composition of wood. The regression analysis and typical correlation analysis of 26 characters, such as growth, material property, physical mechanics, chemical composition and so on, found that the main influencing factors that affect the three factors of the tree trunk deformation are the height of the tree, the volume of the material, the diameter of the breast, the thickness of the bark, the fiber length, Dry shrinkage rate, tensile strength, tensile elastic modulus and total cellulose content. in which, the growth characteristics of the Eucalyptus globulus which have the greatest influence on the wind resistance of the eucalyptus are the tree height, the material product, the DBH, the second is the fiber length, the tensile strength, and the genetic diversity analysis and test material is 109 samples of the eight eucalyptus groups, A total of 107 SSR loci covering the whole genome were identified.55 neutral genomic SSRs revealed a high diversity of Eucalyptus globulus, and a total of 1116 alleles were detected, with an average of 20.3 alleles per point, a PIC of 0.35508-0.9489, an average of 0.771, and a site with a higher polymorphism. The mean value of HE is 0.718, which indicates that the height of Eucalyptus is heterozygous, the average of Fst is 0.066, the genetic differentiation among the groups is not significant, and the variance component of the inter-population variance in the molecular analysis of variance is only 1%, indicating that the genetic variation is mainly present in the population. The genetic background of the subpopulation in Australia was found to be different from the genetic distance analysis of Nei (1972). The population of Eucalyptus grandis has rich genetic diversity, and can be used for the breeding of wind-resistant strains and the genetic improvement of the gene library. The SSR-associated analysis of the resistance-related traits of Eucalyptus urophylla was obtained by using 107 SSR markers with a large difference in the medium-position frequency of two types of gene pools. There was a significant linkage with 26 SSR loci (P0.05), which could explain the growth and resistance to the phenotypic variation of 9.26% ~ 71.14%. Among the 26 associated sites, the rate of the ECeSSR235 site to the wind-hazard index was 71.14%, the lowest in the ECeSSR570 site was 9.26%, and the average interpretation rate was 36.27%. The site ECeSSR332 was associated with four traits such as tree height, DBH, volume and wind-damage index, and the interpretation rate of the four traits was relatively high, over 60%. The site EUCSSR484, EUCSSR352, EUCSSR570 and EUCSSR422 were associated with two traits, such as the high tree height and the wind hazard index, and the EUCSSR489 and EUCSSR114 sites were associated with 2 traits such as tree height and volume.
【學(xué)位授予單位】：中國林業(yè)科學(xué)研究院
【學(xué)位級(jí)別】：博士
【學(xué)位授予年份】：2017
【分類號(hào)】：S792.39
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本文編號(hào)：2511907
文內(nèi)圖片：
圖片說明：09份赤桉種質(zhì)資源群體結(jié)構(gòu)圖
[Abstract]:The frequent occurrence of typhoon and wind damage in the coastal areas of the south of China have seriously affected the ecological security of the coastal areas and the development of the related industries, in particular, it poses a great threat to the growth of the trees, and becomes one of the important factors to restrict the sustainable development of the eucalyptus plantation. The breeding of the good wind-resistant strain is an important measure to ensure the sustainable development of the eucalyptus goods in the coastal area of China. Eucalyptus camaldulensis is an important genetic material for the improvement of eucalyptus tree species in China. In this paper,26 characters, such as growth, material property, physical mechanics, chemical composition and so on, were investigated by using 114 red eucalyptus family (half-siblings), and the main influencing factors that affected the wind resistance of the eucalyptus were found through the analysis of the genetic variation and the correlation analysis. The wind and heavy load deformation and the wind-resistant main influence factors are analyzed by the wind damage simulation (Rtree experiment), and the wind resistance characteristics of the Eucalyptus urophylla family are evaluated, and the main influence factors affecting the wind resistance of the Eucalyptus urophylla are obtained, and the excellent anti-wind family is selected. The wind resistance of Eucalyptus is caused by a number of complex factors. This study is based on the main influencing factors that affect the wind resistance of the eucalyptus, and uses the SSR molecular marker technology to carry out the correlation analysis. The correlation analysis of the wind resistance of the eucalyptus is carried out, and the genetic basis of the wind resistance of the eucalyptus is analyzed. And the excellent SSR locus is excavated. This study will help to establish a molecular marker-assisted selection system for wind-resistant breeding of eucalyptus, laying the foundation for exploring and using the important anti-wind gene resources of the eucalypt, and providing a theoretical basis for the follow-up research. The main results of the study are as follows:1. There are abundant genetic variation among the growth characters of the family of Eucalyptus, and the heritability of the family is higher than the heritability of the single plant, and the selection potential of the family is larger. There are abundant genetic variation among the characters of the early Eucalyptus (1-year-old), which has a strong family and single-plant selection potential in the family. The genetic correlation coefficient between the wind-damage index and the growth character of the family is the largest, the resistance to wind resistance and the size of the tree is not strong, and the regression equation also shows that the resistance to wind resistance is not related to the growth character of the family, and the faster the tree plant grows (i.e. the larger the volume product), The results showed that the average wind-damage index was below grade 2, and the wind-hazard index of the forest was negatively correlated with the tree size (tree height and single-plant volume). The genetic control level of some characters was relatively high. The correlation of the wind-damage grade of the 3-4-year-old Eucalyptus family with the tree height, the crown width, the volume of the material and the Pearson correlation of the bark thickness was negatively correlated. It is indicated that the higher the tree, the greater the volume of the material, and the wind resistance of the 3-4-year-old Eucalyptus urophylla family shows that the wind resistance of the eucalyptus is related to the height of the tree, the diameter of the breast and the volume of the tree. There was no significant difference in the inter-family and family and site interactions among the different families, and the wood density of the 40-month-old Eucalyptus grandis belongs to the small grade, which belongs to the light wood. According to the physical and mechanical classification standard of the wood, the dry shrinkage rate of the eucalyptus is grade III, is easy to crack, the dry shrinkage value of the difference between the bottom of the trunk and the middle part of the trunk is relatively large, and the difference in the dry shrinkage of the tip part is large, so that the bottom is more prone to cracking and deformation. The difference of the mechanical properties (bending strength, flexural modulus, the shear strength and the compressive strength) of the different parts of Eucalyptus grandis is very significant, the difference of the flexural modulus of the different parts is very significant, and the difference of the other indexes is not significant. The study found that the size of trees (height, DBH, volume of wood) was the main factor that affected the wind resistance of the eucalyptus, followed by a material factor. The degree of wind damage of a single plant is closely related to the size of the vertical wood, and also has a strong correlation with the wood properties such as the fiber width, the Pilodyn value and the stress wave value. The correlation between the mechanical properties such as the basic density of the wood and the dry shrinkage, the physical mechanics (bending strength, the flexural modulus of bending, the shear strength of the line, the compressive strength of the smooth grain, etc.) and the anti-wind resistance of the eucalyptus urophylla can be seen to show that the correlation between the two is not significant, and the wind-hazard index is similar to that of the lignin and the heddle, The correlation between cellulose and hemicellulose was not significant (P0.05). From the correlation analysis, it was found that the anti-wind property of Eucalyptus globulus was not related to the physical and chemical composition of wood. The regression analysis and typical correlation analysis of 26 characters, such as growth, material property, physical mechanics, chemical composition and so on, found that the main influencing factors that affect the three factors of the tree trunk deformation are the height of the tree, the volume of the material, the diameter of the breast, the thickness of the bark, the fiber length, Dry shrinkage rate, tensile strength, tensile elastic modulus and total cellulose content. in which, the growth characteristics of the Eucalyptus globulus which have the greatest influence on the wind resistance of the eucalyptus are the tree height, the material product, the DBH, the second is the fiber length, the tensile strength, and the genetic diversity analysis and test material is 109 samples of the eight eucalyptus groups, A total of 107 SSR loci covering the whole genome were identified.55 neutral genomic SSRs revealed a high diversity of Eucalyptus globulus, and a total of 1116 alleles were detected, with an average of 20.3 alleles per point, a PIC of 0.35508-0.9489, an average of 0.771, and a site with a higher polymorphism. The mean value of HE is 0.718, which indicates that the height of Eucalyptus is heterozygous, the average of Fst is 0.066, the genetic differentiation among the groups is not significant, and the variance component of the inter-population variance in the molecular analysis of variance is only 1%, indicating that the genetic variation is mainly present in the population. The genetic background of the subpopulation in Australia was found to be different from the genetic distance analysis of Nei (1972). The population of Eucalyptus grandis has rich genetic diversity, and can be used for the breeding of wind-resistant strains and the genetic improvement of the gene library. The SSR-associated analysis of the resistance-related traits of Eucalyptus urophylla was obtained by using 107 SSR markers with a large difference in the medium-position frequency of two types of gene pools. There was a significant linkage with 26 SSR loci (P0.05), which could explain the growth and resistance to the phenotypic variation of 9.26% ~ 71.14%. Among the 26 associated sites, the rate of the ECeSSR235 site to the wind-hazard index was 71.14%, the lowest in the ECeSSR570 site was 9.26%, and the average interpretation rate was 36.27%. The site ECeSSR332 was associated with four traits such as tree height, DBH, volume and wind-damage index, and the interpretation rate of the four traits was relatively high, over 60%. The site EUCSSR484, EUCSSR352, EUCSSR570 and EUCSSR422 were associated with two traits, such as the high tree height and the wind hazard index, and the EUCSSR489 and EUCSSR114 sites were associated with 2 traits such as tree height and volume.
【學(xué)位授予單位】：中國林業(yè)科學(xué)研究院
【學(xué)位級(jí)別】：博士
【學(xué)位授予年份】：2017
【分類號(hào)】：S792.39
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本文編號(hào)：2511907
本文鏈接：http://sikaile.net/shoufeilunwen/nykjbs/2511907.html
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