【摘要】：紅豆樹(shù)(Ormosia hosiei)為國(guó)家二級(jí)重點(diǎn)保護(hù)植物,自然分布于陜西南部和江蘇、浙江、江西、福建、湖北、四川、重慶、貴州等省(市),呈星散間斷或小塊狀生于溪邊、山谷常綠闊葉林中。紅豆樹(shù)材質(zhì)優(yōu)良,有光澤,其紋理美觀(guān),是制作工藝雕刻、裝飾鑲嵌等的上等用材。由于紅豆樹(shù)具有較高的經(jīng)濟(jì)和人文價(jià)值,其原有天然資源被遭到嚴(yán)重的人為破壞。紅豆樹(shù)現(xiàn)有天然居群較小,自然更新困難,已經(jīng)處于瀕危狀態(tài),但目前對(duì)其遺傳背景了解較少。本研究在已有研究基礎(chǔ)上,首先,開(kāi)發(fā)適用于紅豆樹(shù)遺傳多樣性分析的SSR標(biāo)記,系統(tǒng)研究不同小流域紅豆樹(shù)天然居群的遺傳多樣性和遺傳分化;其次,通過(guò)研究天然居群和個(gè)體的交配系統(tǒng)及親子代遺傳差異,闡明紅豆樹(shù)天然居群高遺傳多樣性的維持機(jī)制;最后,開(kāi)展優(yōu)樹(shù)子代苗期測(cè)定,研究苗期生長(zhǎng)和形態(tài)的家系變異及所受遺傳控制,揭示子代遺傳多樣性對(duì)子代生長(zhǎng)的影響,以期為紅豆樹(shù)遺傳保育策略制定及資源有效利用提供理論依據(jù)。主要研究結(jié)果如下:1、利用SLAF-seq(Specific-locus amplified fragment sequencing)技術(shù)對(duì)紅豆樹(shù)基因組進(jìn)行簡(jiǎn)化測(cè)序,獲得6 426 462 reads和16 653個(gè)多態(tài)性SLAF標(biāo)簽,含有17 868個(gè)SSR位點(diǎn),平均每6.98kb分布1個(gè)SSR位點(diǎn)。不同核苷酸重復(fù)類(lèi)型的基元類(lèi)型數(shù)量及SSR位點(diǎn)數(shù)量間差異較大,除單核苷酸外,二、三核苷酸重復(fù)類(lèi)型數(shù)量較多,分別占總SSR位點(diǎn)的17.15%和15.02%,其中AT/TA和GAA/TTC基元重復(fù)數(shù)量最多,分別為1 090個(gè)(43.1%)和349個(gè)(15.2%)。在獲取的2 817對(duì)引物中,以二、三核苷酸類(lèi)型最多,兩者所占比例高達(dá)94.8%。從設(shè)計(jì)合成的70對(duì)SSR引物中,篩選出12對(duì)多態(tài)性高、特異性好的SSR引物,多態(tài)位點(diǎn)百分率為17.14%。這為紅豆樹(shù)遺傳多樣性、交配系統(tǒng)分析及分子標(biāo)記輔助育種奠定了基礎(chǔ)。2、利用開(kāi)發(fā)的SSR引物對(duì)分布在江西省資溪縣瀘溪河(LXH)、福建省柘榮縣茜洋溪(XYX)和浙江省龍泉市甌江上游(OJ)3個(gè)小流域紅豆樹(shù)天然居群的遺傳分析發(fā)現(xiàn),不同小流域紅豆樹(shù)天然居群皆維持較高遺傳多樣性水平(HE0.720)。不同流域紅豆樹(shù)天然居群遺傳多樣性的高低順序:OJ(HE=0.835)、XYX(HE=0.829)、LXH(HE=0.796);西溪支流內(nèi)中游的(福建富溪)FJFX居群遺傳多樣性明顯高于其上游福建東源(FJDY)和下游福建宅中居群(FJZZ)。遺傳變異主要存在于流域內(nèi)(92.73%)和居群內(nèi)(89.66%),流域間或流域內(nèi)居群間的遺傳分化均屬于中等程度(Fst分別為0.070和0.103)�；谶z傳距離的聚類(lèi)分析和Structure分組分析,均將紅豆樹(shù)3個(gè)小流域的9個(gè)天然居群歸為2大群組,其中LXH和XYX流域歸為第一群組,而OJ流域?yàn)榈诙航M。遺傳距離和地理距離之間無(wú)顯著線(xiàn)性相關(guān)。據(jù)此,推測(cè)紅豆樹(shù)片段化時(shí)間較短,未對(duì)其遺傳多樣性產(chǎn)生較大影響。3、在對(duì)紅豆樹(shù)天然居群遺傳多樣性分析基礎(chǔ)上,選取江西馬頭山(JXMTS)、浙江八都(ZJBD)和浙江錦溪-1(ZJJX-1)3個(gè)典型的天然居群,進(jìn)一步對(duì)其交配系統(tǒng)進(jìn)行分析,結(jié)果表明,成株居群和子代群體均保持了較高的遺傳多樣性,不同居群及其子代的遺傳多樣性略有差異,其中JXMTS和ZJBD居群遺傳多樣性高于ZJJX-1居群。子代群體固定指數(shù)(F)均明顯低于成株居群,說(shuō)明子代群體中實(shí)際觀(guān)察到的純合單株所占比例要高于親本。紅豆樹(shù)的異交率較高(tm=0.884),存在一定程度的雙親近交現(xiàn)象(tm ts=0.078),且花粉供體數(shù)目較少(2.096)。JXMTS(tm=1.000)的異交率高于ZJBD居群(tm=0.824),而大的居群有效花粉供體數(shù)相對(duì)較少。取樣8個(gè)家系的多位點(diǎn)異交率(tm)變化范圍為0.772~1.000,絕大多數(shù)子代家系均存在一定程度的雙親近交現(xiàn)象,個(gè)別家系有少量自交。這些結(jié)果說(shuō)明創(chuàng)造條件擴(kuò)大居群規(guī)模對(duì)紅豆樹(shù)天然居群的就地保護(hù)非常重要,還應(yīng)防止邊緣或孤立個(gè)體的人為破壞而導(dǎo)致基因流受阻。4、利用在浙、閩、贛和川4省初選的76株優(yōu)樹(shù),開(kāi)展紅豆樹(shù)優(yōu)樹(shù)種子及其苗期生長(zhǎng)遺傳變異研究,結(jié)果表明,紅豆樹(shù)優(yōu)樹(shù)種子性狀(種寬、種厚等)及其子代生長(zhǎng)(苗高和地徑)、葉片形態(tài)和根系特征等性狀大多具有顯著或極顯著的家系效應(yīng),高生長(zhǎng)節(jié)律參數(shù)在家系間均達(dá)到極顯著水平。相關(guān)分析結(jié)果顯示,苗高、地徑與種厚、百粒重均呈極顯著正相關(guān),較之于苗高,地徑與葉片、根系特征等性狀的相關(guān)關(guān)系更強(qiáng)(r=0.349~0.608);苗高與線(xiàn)性生長(zhǎng)始期、各生長(zhǎng)參數(shù)(最大線(xiàn)性生長(zhǎng)速度、線(xiàn)性生長(zhǎng)速度和線(xiàn)性生長(zhǎng)量)均呈極顯著正相關(guān),而與線(xiàn)性生長(zhǎng)期呈微弱負(fù)相關(guān),說(shuō)明相對(duì)較晚進(jìn)入速生期的紅豆樹(shù)家系,憑借較快的生長(zhǎng)速率更能優(yōu)先占據(jù)生長(zhǎng)空間而獲得較高的生長(zhǎng)量。各生長(zhǎng)性狀家系遺傳力水平較高,受中等或較強(qiáng)的遺傳控制(hf2=0.479~0.854),并以苗高作為主要性狀,同時(shí)兼顧地徑,從76個(gè)優(yōu)樹(shù)家系中初選出20個(gè)優(yōu)良家系�？偟膩�(lái)說(shuō),種子性狀、葉片及生長(zhǎng)節(jié)律參數(shù)等對(duì)苗木生長(zhǎng)影響顯著,為今后紅豆樹(shù)優(yōu)良家系選擇提供參考。5、選擇26個(gè)代表性紅豆樹(shù)優(yōu)樹(shù)自由授粉子代群體,進(jìn)行子代群體遺傳多樣性評(píng)價(jià)及其對(duì)苗木生長(zhǎng)影響的研究,分析發(fā)現(xiàn),紅豆樹(shù)優(yōu)樹(shù)子代群體亦具有較高的遺傳多樣性,有效等位基因數(shù)為7.766個(gè),觀(guān)測(cè)雜合度(HO)和期望雜合度(HE)分別為0.469和0.865。除SSR8外,其余位點(diǎn)的觀(guān)測(cè)雜合度均小于期望雜合度,表明子代群體絕大多數(shù)位點(diǎn)處于雜合子缺失狀態(tài)。不同家系紅豆樹(shù)的遺傳多樣性有所差異,12號(hào)家系的遺傳多樣性水平最高,8號(hào)家系最低。通過(guò)比較發(fā)現(xiàn),居群子代的遺傳多樣性顯著或極顯著高于孤立木子代。F統(tǒng)計(jì)量和分子方差分析(AMOVA)均表明,紅豆樹(shù)優(yōu)樹(shù)子代群體的遺傳變異主要存在于家系內(nèi),家系間的遺傳分化較小。相關(guān)性分析發(fā)現(xiàn),子代遺傳多樣性參數(shù)與種子性狀、子代播種后7個(gè)月的苗高呈顯著正相關(guān)(r=0.378~0.527)。說(shuō)明較大的紅豆樹(shù)天然居群在子代遺傳多樣性維系中起了非常重要作用,子代遺傳多樣性顯著影響苗木后期生長(zhǎng),這為后期優(yōu)良家系選擇提供了理論依據(jù)。
[Abstract]:Ormosia hosiei is a national secondary key protection plant, which is naturally distributed in the south of Shaanxi and the provinces (cities) of Jiangsu, Zhejiang, Jiangxi, Fujian, Hubei, Sichuan, Chongqing, Guizhou and other provinces (cities). The red bean tree has the advantages of excellent material and luster, and the texture is beautiful, and the red bean tree has the advantages of manufacturing process engraving, decoration and inlaying, and the like. Because the red bean tree has higher economic and cultural value, the original natural resources of the red bean tree are destroyed artificially. The existing natural living group of the red bean tree is small, the natural updating is difficult, and the red bean tree is in an endangered state, but the genetic background of the red bean tree is less. Based on the existing research, the genetic diversity and genetic differentiation of the natural population of the red bean trees in different small watersheds were studied by using SSR markers suitable for the analysis of the genetic diversity of the red bean trees, and secondly, By studying the genetic difference of the mating system and the progeny of the natural population and the individual, the maintenance mechanism of the high genetic diversity of the natural habitat of the red bean tree was clarified, and the family variation and the genetic control of the growth and the morphology in the seedling stage were studied. The influence of the genetic diversity of the offspring on the growth of the offspring is revealed, with a view to providing the theoretical basis for the development of the genetic conservation strategy and the effective utilization of the resources of the red bean trees. The main results of this study were as follows:1. Using the SLAF-seq (Specific-location) technique to simplify the sequencing of the genome of the red bean trees to obtain the 6426 462-reads and 16,653 polymorphic SLAF tags, including 17,868 SSR sites, with an average of 1 SSR locus per 6.98 kb. The number of the primitive types and the number of SSR loci of the different types of nucleonic acid repeats is large, and the number of the repeat types of the two and the three-core acid is more than that of the single-core acid, which accounts for 17.15% and 15.02% of the total SSR locus, respectively, wherein the repeat number of the AT/ TA and the GAA/ TTC elements is the most, 1,090 (43.1 per cent) and 349 (15.2 per cent), respectively. Among the 2,817 pairs of primers, the most of the two, three, and third, the proportion of the two were as high as 94.8%. 12 pairs of SSR primers with high polymorphism and good specificity were selected from 70 SSR primers which were designed and synthesized. The percentage of polymorphic sites was 17.14%. This provides a basis for the genetic diversity, mating system analysis and molecular marker-assisted breeding of the red bean tree. The genetic analysis of the natural population of the red bean trees in the three small watershed of the three small watershed of the Qiangxi (XYX) and the upstream (OJ) of the Longquan city of the province of Fujian Province found that the natural population of the red beans in the different small watershed maintained a higher level of genetic diversity (HE0.720). The genetic diversity of the natural populations of the red beans in different watersheds is: OJ (HE = 0.835), XYX (HE = 0.829), LXH (HE = 0.796); the genetic diversity of the FJFX population in the middle reaches of the Xixi substream is significantly higher than that of the upstream Fujian Dongyuan (FJDY) and the downstream Fujian house (FJZZ). The genetic variation is mainly in the basin (92.73%) and within the population (89.66%), and the genetic differentiation among the interbasin or interbasin populations is of moderate degree (Fst is 0.070 and 0.103, respectively). Based on the cluster analysis and the structure analysis of the genetic distance, the nine natural groups in the 3 small watershed of the red bean tree are classified as two groups, and the LXH and XYX basins are classified as the first group, and the OJ drainage area is the second group. There was no significant linear correlation between the genetic distance and the geographic distance. Based on the analysis of the genetic diversity of the natural population of the red beans, the three typical natural populations of the Maheadshan (JXMTS) of Jiangxi, eight of Zhejiang (ZJBD) and Jinxi-1 (ZJJX-1) of Zhejiang Province were selected. The results showed that the genetic diversity of JXMTS and ZJBD populations was higher than that of ZJJX-1 populations. The fixed index (F) of the progeny population was significantly lower than that of the parent population, indicating that the proportion of the homozygous single plant observed in the progeny population was higher than that of the parent. The heterogenic rate of red bean trees was higher (tm = 0.884), and there was a certain degree of parental close-crossing (tm ts = 0.078), and the number of pollen donors was less (2.096). The cross-crossing rate of JXMTS (tm = 1.000) was higher than that of the ZJBD population (tm = 0.824), while the number of effective pollen donors in large populations was relatively small. The variation range of the multi-locus of 8 families was 0.772 ~ 1.000, and the most of the children's families had a certain degree of close-crossing, and the individual families had a small number of selfing. These results indicate that the condition of creating conditions to enlarge the population size is very important to the local protection of the natural population of the red bean trees, and the human-made destruction of the edge or the isolated individual should also be prevented, and the flow of the gene can be prevented. The results of the study on the genetic variation of the seed and its seedling at the seedling stage show that the characteristics of the sub-traits (seed width, seed thickness, etc.) and its progeny growth (high seedling height and ground diameter), leaf shape and root system characteristics of the superior tree species have significant or very significant family effects. The high growth rhythm parameters reached a very significant level at home. The results of correlation analysis showed that the seedling height, the ground diameter and the seed thickness and the weight average of the seed were positively correlated, and the correlation between the ground diameter and the leaf and root system characteristics was stronger (r = 0.349-0.608), and the growth parameters (maximum linear growth rate) were higher in the seedling height and the linear growth stage, and the growth parameters (maximum linear growth rate). The linear growth rate and the linear growth rate were positively correlated with a slight negative correlation with the linear growth phase, indicating that the red bean tree family was relatively late to enter the fast growing stage, and the growth space was more preferentially occupied by the faster growth rate to obtain higher growth. The heritability of each growth trait family was high, and it was controlled by medium or strong genetic control (hf2 = 0.479-0.854), and with high seedling height as the main character, and taking into account the ground diameter,20 excellent families were selected from the 76 excellent tree families. in general, that effect of the seed character, the leaf and the growth rhythm parameter on the growth of the nursery stock is significant, provide reference for the good family selection of the red bean tree in the future, and 5, select the free-pollinated progeny group of the 26 representative red bean trees, The genetic diversity of the progeny population and its effect on the growth of the nursery stock were studied. The results showed that there were also high genetic diversity, the number of effective alleles was 7.766, the observed heterozygosity (HO) and the expected heterozygosity (HE) were 0.469 and 0.865, respectively. In addition to the SSR8, the observed heterozygosity of the remaining sites was less than the expected heterozygosity, indicating that the majority of the sites of the progeny population were in the heterozygote-deleted state. The genetic diversity of the red bean trees in different families is different, and the genetic diversity of the family in No.12 family is the highest and the family of the No.8 families is the lowest. It was found that the genetic diversity of the children in the population group was significantly or significantly higher than that of the isolated wood. F-statistic and variance analysis of variance (AMOVA) showed that the genetic variation of the progenies of the superior tree of the red bean was mainly found in the family, and the genetic differentiation among the families was small. The correlation analysis showed that the genetic diversity of the progeny was positively correlated with the seed character and the seedling height of 7 months after the seed sowing (r = 0.378-0.527). It is indicated that the natural population of red bean trees plays an important role in the maintenance of the genetic diversity of the progeny, and the genetic diversity of the progeny significantly influences the later growth of the nursery stock, which provides a theoretical basis for the good family selection in the later stage.
【學(xué)位授予單位】：中國(guó)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(lèi)號(hào)】：S792.99

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