结缕草属（Zoysia Willd.）植物种质资源多样性研究
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摘要
结缕草属(Zoysia Willd.)草坪草具有低维护、耐践踏、耐干旱、耐瘠薄、抗病虫害以及耐盐碱等特性,是世界著名的环保型暖季草坪草。结缕草属植物广泛分布于太平洋西岸和北岸沿海地区,在非洲和澳洲等温暖地区也有分布。
本研究以课题组搜集的结缕草属种质资源为试验材料,对中国结缕草属的5种1变种代表性种源的重要坪用性状、物候期、种子产量及相关性状、营养繁殖速度进行了研究；应用改良电导法对该属5种1变种97份种源的抗寒性进行了初步鉴定分析,在此基础上应用生长恢复法对其中不同抗寒性的代表种源的匍匐茎和根状茎的抗寒性进行了研究,分析了不同器官在遭受低温胁迫时的抗寒性之间的关系,随后挑选了两个抗寒性极端的种源,对叶片、匍匐茎和根状茎三个不同器官在低温胁迫后的可溶性蛋白进行SDS-PAGE电泳分析；利用形态学标记、等位酶标记、SSR分子标记三种不同水平上的遗传标记技术结合地理分布对结缕草属植物的9种1变种59份种源的遗传变异进行分析,比较了种内遗传分化大小,分析了这些种在进化过程中的亲缘关系。上述研究将为结缕草属植物种质资源开发利用提供重要的科学依据。
全文主要结论如下：
1中国结缕草属(Zoysia Willd.)植物外部形状变异分析
对中国结缕草属5种1变种共计124份种源的14个外部性状的变异及其规律进行了研究。结果表明：①中国结缕草属植物的14个性状的变异都很大,变异系数都超过了10%。②结缕草属植物外部性状间存在一定的相关性,如密度与除小穗长以外其他所有的性状间都呈显著的负相关。③结缕草和中华结缕草的某些外部性状与经、纬度之间存在一定的相关性。④124份结缕草属种源划分为4组,细叶型结缕草、多花粗叶型结缕草、少花粗叶型结缕草及Z099单独为一组。
2中国结缕草属(Zoysia Willd.)植物物候期变异分析
系统观测分析了中国结缕草属5种1变种共109份种源的物候期后,结果表明,在南京地区,中国结缕草属植物物候期存在不同程度的变异,其中盛花期和枯黄期的变异最大,青绿期变异范围为215～277d；孕穗初期的早晚与初花期、盛花期、结实期和枯黄期的早晚及青绿期的长短都存在显著正相关,返青期的早晚与青绿期的长短呈极显著负相关,而枯黄期的早晚与青绿期的长短则呈极显著正相关,返青期的早晚与枯黄期的早晚不存在相关关系；结缕草属植物的返青期、孕穗初期、初花期、盛花期及枯黄期随纬度增加而显著提前。相对的质地粗的种源的物候期比质地细的种源要早。
3中国结缕草属(Zoysia Willd.)植物繁殖性能研究
在对中国结缕草属植物结缕草(Z.japonica Steud.)和中华结缕草(Z.sinica Hance.)的40份种源的匍匐茎营养繁殖速度、种子产量及种子产量组分调查分析后,发现,在南京地区：(1)匍匐茎总长度和地上部变异系数(CV)分别是69.57%和32.45%,变异很大；(2)匍匐茎总长度和地上部生物量两个指标间呈极显著的正相关,相关系数为0.676；(3)花序密度、单位花序种子数、百粒重以及种子产量方面存在显著差异,其中种子产量变异程度最高,CV达到102.51%；(4)单位花序种子数、百粒重与种子产量间正相关,种子产量的3个组分相互作用后与种子产量复相关；(5)通径分析表明,单位花序种子数、百粒重两个组分对种子产量的贡献很大,而花序密度对种子产量的贡献较小(6)地上部生物量与花序密度呈显著负相关；(7)对匍匐茎总长度、种子产量多重比较后发现Z015、Z013、Z056等6份种源营养生长速度快,Z016、Z052、Z061等9份种源种子产量高,Z012、Z063等6份种源,营养生长速度快且种子产量也高,而Z111、Z018等5份种源,既没有高的种子产量,其营养生长速度又慢。
4结缕草属(Zoysia Willd.)植物抗寒性研究
4.1结缕草属植物抗寒性鉴定
采用改良电导法初步鉴定了结缕草属植物5种1变种97份种源叶片抗寒性,在此基础上,选取其中15份抗寒性有差异的种源的匍匐茎和根状茎进行生长恢复试验。结果表明：(1)以叶片半致死温度(LT50)为评价指标,结缕草属植物的LT50的范围是-1.9℃～-10.4℃,变异系数为27.79%；结缕草属植物不同种的LT50从低到高依次为：结缕草<中华结缕草<沟叶结缕草(Z.matrella (L.) Merr.)<细叶结缕草(Ztenuifolia Thiele.)<长花结缕草(Z.sinica var longiflora Ohwi.)<大穗结缕草(Zmacrostachya Franchet et Sav.);(2)结缕草、中华结缕草和沟叶结缕草三个种种内LT50的变异都很大,变异系数(CV)都高于20%。(3)不同季节的匍匐茎和根状茎的LT50间存在显著的正相关。(4)不同器官的LT50间的R均达到了显著水平。
4.2结缕草冷驯化条件下可溶性蛋白变化分析
通过对极端抗寒性的两个结缕草种源Z125(弱)和Z141(强)的叶片、匍匐茎和根状茎三个器官在正常生长条件(28℃/22℃,白天/黑夜,8h光照)与模拟低温驯化(8℃/2℃,白天/黑夜,光照8h)后的可溶性蛋白的SDS-PAGE电泳分析,发现,在模拟低温处理后2周,抗寒性差的Z125的可溶性蛋白合成发生较大的变化,根状茎比对照多了44KD的一条带,叶片比对照多了66KD左右的带,而抗寒性强的Z141仅在根状茎中比对照多了一条50KD的带,但少了一条40KD左右的带；处理4周后,Z141的匍匐茎和根状茎的可溶性蛋白都比对照多了两条带,大小为60KD左右,其叶片的可溶性蛋白电泳条带比未驯化的少了2条带(分别是60KD和20KD左右),而Z125仅根状茎可溶性蛋白比对照多了两条带,其多肽大小约为50KD。
5结缕草属(Zoysia Willd.)植物遗传变异和种间关系
5.1形态学研究
通过对结缕草属植物9种1变种59份种源的23个外部性状的观测和分析,得出结论：①沟叶结缕种内各种源间在很多性状上都存在很大的变异,结缕草和中华结缕草的种内变异也较大,次于沟叶结缕草,并且两者的种内变异大小较接近。朝鲜结缕草(Z. koreana Mez)种内变异小；②这些种之间根据遗传距离远近可以分成三大类：第一类为结缕草、中华结缕草、朝鲜结缕草先聚在一起之后再和大穗结缕草聚成一大类；第二类为沟叶结缕草和细叶结缕草先聚为一小类,长花结缕草和小结缕草(Zminima Zotov)聚为另一小类,然后这些种聚成一大类。大花结缕草(Z. macrantha Desvaux)独自成一集群为第三类；③主成分分析结果表明：叶片宽度,匍匐茎长度、直径,每穗小穗数,小穗柄长度,小穗长度,生殖枝高度,叶舌的颜色及有无毛这9个性状对结缕草属植物种间亲缘关系的研究具有重要的意义。
5.2等位酶研究
分析结缕草属植物9种1变种59份种源的苹果酸脱氢酶(MDH),酯酶(EST)及过氧化物酶(POD)三种等位酶特征,研究其种间亲缘关系,结果表明：①种内遗传分化大小依次为：沟叶结缕草>结缕草>中华结缕草>朝鲜结缕草；②聚类结果显示：结缕草、中华结缕草、长花结缕草、大穗结缕草聚为第一类；沟叶结缕草、细叶结缕草先聚为一类,然后小结缕草、太平洋结缕草(Z. pacifica Hotta)、大花结缕草也聚到这一类；而朝鲜结缕草自成一类。
5.3SSR分子标记研究
利用SSR分子标记对结缕草属植物9种1变种59份种源进行了种间关系分析,结果表明：①种内遗传分化大小依次为：结缕草、中华结缕草>朝鲜结缕草>沟叶结缕草；②聚类结果显示：结缕草、中华结缕草、朝鲜结缕草和大穗结缕草聚为一类；沟叶结缕草、小结缕草和细叶结缕草三个种聚为一类；大花结缕草、长花结缕草各自为一类。
6中国结缕草属种质资源综合评价
应用叶色、密度、叶宽、均一性、青绿期、抗寒性、锈病抗性7个指标,对中国结缕草属植物的74份种源在南京地区的坪用价值进行评价,结果表明：Z014、Z075、Z008、Z142、Z042、Z005、Z044、Z101、Z053、Z050等种源叶色较深而密度较高,青绿期较长但抗寒性一般,综合坪用质量高；而Z118、Z081、Z010、Z115、Z104、Z103等种源叶色黄绿,青绿期较短,抗寒性较差,锈病抗性差,综合坪用质量低。其他54份种源由于2个或3个指标得分较低,综合坪用质量中等。
Zoysia grasses (Zoysia spp.) are widely used as warm season turf grass indigenous to the countries of the western Pacific Rim. They possess excellent wear tolerance, drought tolerance, disease resistance, salinity tolerance, low fertility tolerance and low maintenance requirements.
Forteen morphological characteristics, phonological phase, seed yield and its component, stolon growth rate of representative accessions of5species and1variety of Zoysia Willd. were observed in this report. The leaf low temperature of97accessions of5species and variety of Zoysia Willd. was evaluated using modified electrolyte leakage method, and15representative accessions were selected based on leaf low temperature result to evaluate the low-temperature tolerance of stolons and rhizomes using survival rate, and then soluble protein of three organs of two extreme low temperature tolerant representative accessions were studied using SDS-PAGE electrophoresis. Morphology, isozyme and SSR genetic markers at three different levels were studied combining with the geographical distribution to study the genetic variation and interspecific relationship of59accessions of Zoysia Willd. belonging to9species and1variety. The study will provide an important scientific basis for the development and untilization of zoysiagrass germplasm resource in China.
The main results were stated as followings:
1. Variation analysis on morphological characteristic of Zoysia Willd. in China
The variations and tendencies of14morphological and their correlationship of124accessions of Zoysia Willd. were studied in the paper. The results showed that the great variation of morphological characteristics were existed among the accessions of Zoysia Willd.with the coefficient of variation (CV) at exceed10%. Correlation analysis indicated obvious correlation were existed among some of14morphological characteristics at significant levels. Some morphological characteristics was correlated siginificantly with latitude or longitude. Four groups were divided based on14morphological characteristics: fine zoysiagrass, inflorescenced coarse zoysiagrass, less-inflorescenece coarse zoysiagrass and Z099.
2. Study on the variation of phenological phase of Zoysia Willd.
The phenological phases of109accessions of Zoysia Willd. including5species and1variety were observed. The results showed that there is difference in the date of phonological phases among different accessions in Nanjing. Variation scopes of blooming stage and wilting stage was biggest, and that of green period was215-277d. The date of initial booting stage has a significantly positive correlation with the date of initial blooming stage, blooming stage, maturation stage and wilting stage and length of green period. The date of wilting stage has a positive correlation with the length of green period, and that of turning-green stage has a highly significant negative correlaion with the length of green period. Regression analysis indicated that turing green, initial booting, initial blooming, blooming and wilting stages of Zoysia Willd. became obviously earlier with latitude increasing. The phonological phase of coarse texture accessions was earier than fine texture accessions relatively.
3. Study on propagation properties of Zoysia Willd. in China
The stolon growth rate, the seed yield and its component of Z. japonica and Z. sinica in China were measured and analyzed. The results showed that in Nanjing:(1) abundant variation was existed in the total stolon length and aboveground biomass of different accessions of zoysiagrass;(2) the total stolon length was positive corrected with the aboveground biomass with correlation coefficient as0.676;(3) abundant variation was existed in the seed yield and its component, in which seed yield had the highest variation with the coefficient vatiation (CV) as102.51%;(4) seed number per spike and the weight per100seeds were positive corrected with seed yield, inflorescence density was not corrected with seed yield directly, but seed yield was correlated with inflorescence density, seed number per spike and the weight per100seeds after they interacted;(5) path analysis showed that seed number per spike and the weight per100seeds contributed the most to the seed yield, inflorescence density contributed few;(6) above ground biomass was negatively correlated siginificantly with inflorescence density;(7) the multiple comparisons analysis of the total stolon length and seed yield revealed the vegetative growth rate of six accession such as Z015,Z013, Z056et al was high, nine accessions such as Z016, Z052and Z061et al have high seed yield, six accessions such as Z012, Z063et al have high seed yield and vegetative growth rate, and the vegetative growth and seed yield of five accessions such Z111, Z018et al was both low.
4. Study on Low Temperature Tolerance of Zoysia Willd.
4.1The assessment of the Low Temperature Tolerance of Zoysia Willd.
The leaf LT50of97accessions of5species and1variety of Zoysia Willd. were evaluated by electrolyte leakage, then stolon and rhizome low temperature of15representative accession were evaluated by survival rate. The leaf LT50of Zoysia Willd was from-1.9℃to-10.4℃with coefficient of variation (CV) of27.79%, and the LT50of different species of Zoysia Willd were ranked as follows:Z.japonica. 4.2Soluble proteins variation in zoysiagrass after cold-acclimation
Soluble proteins in leaves, stolon and rhizome of Z125(poor cold tolerance) and Z141(strong cold tolerance) in non-acclimation (28℃/22℃, day/night,8h light) and simulated low temperature acclimation(8℃/2℃, day/night,8h light) were studied. SDS-PAGE electrophoresis revealed that after two weeks of cold acclimation, the soluble protein synthesis in Z125had a major change, a new40KD protein components in rhizome and a66KD in leaves in cold-acclimation comparing to the non-acclimation; a more50KD and less40KD protein components changed in rhizome in cold-acclimation Z141comparing to non-acclimation Z141.After4weeks of cold-acclimation, two new protein compositions, about60KD, appeared in cold-acclimated Z141both stolon and rhizome, about60KD and20KD soluble protein compositions were disappeared in cold-acclimated Z141leaf. And only2new soluble protein compositions,about50KD, appeared in cold-acclimated Z125rhizome.
5. Study on Genetic Variation and Interspecific Genetic Relationship of Zoysia Willd.
5.1The morphological markers
With observing and measuring23morphological characteristics on59accessions of Zoysia Willd., the results showed as:The accessions of Z.matrella had greatest variation on most characters, Z.japonica and Z.sinica had great intraspecies differentiation but smaller than Z.matrella, and the degree of genetic diversity within species of these two were very close. The accessions of Z.koreana Mez. had little differentiation on many characters respectively, and the genetic diversity of Z.koreana were smaller;②The cluster analyzing results show:All these species can be classified by genetic distance to be3classes:the first class included of Z. japonica and Z sinica, Z. koreana and Z. macrostachy., the second class included Z. tenuifolia, Z. matrella, Z. minima Zotov. and Z.sinica var. longiflora, Z. macrantha was clustered to be a group itself;③The principal components analyzing results showed that the width of blades, the length and diameter of stolons, the spikelets number of one spike, the length of spikelets and the pedicels, the height of procreant culms, the ligule colour, whether ligule with hairs or not, these features are important in researching of the genetic relationship of species.
5.2Isozyme study
Three isozyme systems (MDH、EST、POD) were conducted on59accessions of Zoysia Willd. to study the genetic relationship of species. The results show as follows:①the genetic diversity was ranked as:Z.matrella> Z. japonica> Z.sinica> Z.koreana.;②the clustering analysis results show:Z. japonica and Z. sinica> Z. sinica var. longiflora and Z.macrostachya were clustered together; Z.matrella、Z. tenuifolia were clustered to be a group, then Z.minima、Z. pacifica、Z. macrantha were also clustered to this group; Z.koreana was clustered to be a group by itself.
5.3SSR analysis
SSR markers were used to study the genetic relationship of accessions of Zoysia Willd..①The genetic diversity within species ranked as:Z.japonica, Z. sinica> Z. koreana> Z. matrella;②Z.japonica was clustered together with Z. sinica, Z. macros tachya and Z koreana; Z. matrella was clustered together with Z. tenuifolia and Z. minima; Z. macrantha and Z.sinica var. longiflora was a group by itself respectively.
6. The evaluation of turf value of Zoysia Willd. germplasm in China
The turf value of74accession of Zoysia Willd.in China were evaluated based on leaf color, density, leaf width, uniformity, green period, low temperature tolerance and rust resistance index. The result showed that ten accessions including Z014、Z075、Z008、Z142、 Z042、Z005、Z044、Z101、Z053and Z050were dark-green, dense, long green-period and common low temperanture tolerance, and the turf quality of these accessions were very high. Ten accessions including Z118、Z081、Z010、Z115、Z104、and Z103et al were yellow-green, short green-period and low tolerancc to low temperature and rust, the turf quality of these ten accessions was very low. The the turf value of other54accessions were intermediated due to two or three poor index.
本研究以课题组搜集的结缕草属种质资源为试验材料,对中国结缕草属的5种1变种代表性种源的重要坪用性状、物候期、种子产量及相关性状、营养繁殖速度进行了研究；应用改良电导法对该属5种1变种97份种源的抗寒性进行了初步鉴定分析,在此基础上应用生长恢复法对其中不同抗寒性的代表种源的匍匐茎和根状茎的抗寒性进行了研究,分析了不同器官在遭受低温胁迫时的抗寒性之间的关系,随后挑选了两个抗寒性极端的种源,对叶片、匍匐茎和根状茎三个不同器官在低温胁迫后的可溶性蛋白进行SDS-PAGE电泳分析；利用形态学标记、等位酶标记、SSR分子标记三种不同水平上的遗传标记技术结合地理分布对结缕草属植物的9种1变种59份种源的遗传变异进行分析,比较了种内遗传分化大小,分析了这些种在进化过程中的亲缘关系。上述研究将为结缕草属植物种质资源开发利用提供重要的科学依据。
全文主要结论如下：
1中国结缕草属(Zoysia Willd.)植物外部形状变异分析
对中国结缕草属5种1变种共计124份种源的14个外部性状的变异及其规律进行了研究。结果表明：①中国结缕草属植物的14个性状的变异都很大,变异系数都超过了10%。②结缕草属植物外部性状间存在一定的相关性,如密度与除小穗长以外其他所有的性状间都呈显著的负相关。③结缕草和中华结缕草的某些外部性状与经、纬度之间存在一定的相关性。④124份结缕草属种源划分为4组,细叶型结缕草、多花粗叶型结缕草、少花粗叶型结缕草及Z099单独为一组。
2中国结缕草属(Zoysia Willd.)植物物候期变异分析
系统观测分析了中国结缕草属5种1变种共109份种源的物候期后,结果表明,在南京地区,中国结缕草属植物物候期存在不同程度的变异,其中盛花期和枯黄期的变异最大,青绿期变异范围为215～277d；孕穗初期的早晚与初花期、盛花期、结实期和枯黄期的早晚及青绿期的长短都存在显著正相关,返青期的早晚与青绿期的长短呈极显著负相关,而枯黄期的早晚与青绿期的长短则呈极显著正相关,返青期的早晚与枯黄期的早晚不存在相关关系；结缕草属植物的返青期、孕穗初期、初花期、盛花期及枯黄期随纬度增加而显著提前。相对的质地粗的种源的物候期比质地细的种源要早。
3中国结缕草属(Zoysia Willd.)植物繁殖性能研究
在对中国结缕草属植物结缕草(Z.japonica Steud.)和中华结缕草(Z.sinica Hance.)的40份种源的匍匐茎营养繁殖速度、种子产量及种子产量组分调查分析后,发现,在南京地区：(1)匍匐茎总长度和地上部变异系数(CV)分别是69.57%和32.45%,变异很大；(2)匍匐茎总长度和地上部生物量两个指标间呈极显著的正相关,相关系数为0.676；(3)花序密度、单位花序种子数、百粒重以及种子产量方面存在显著差异,其中种子产量变异程度最高,CV达到102.51%；(4)单位花序种子数、百粒重与种子产量间正相关,种子产量的3个组分相互作用后与种子产量复相关；(5)通径分析表明,单位花序种子数、百粒重两个组分对种子产量的贡献很大,而花序密度对种子产量的贡献较小(6)地上部生物量与花序密度呈显著负相关；(7)对匍匐茎总长度、种子产量多重比较后发现Z015、Z013、Z056等6份种源营养生长速度快,Z016、Z052、Z061等9份种源种子产量高,Z012、Z063等6份种源,营养生长速度快且种子产量也高,而Z111、Z018等5份种源,既没有高的种子产量,其营养生长速度又慢。
4结缕草属(Zoysia Willd.)植物抗寒性研究
4.1结缕草属植物抗寒性鉴定
采用改良电导法初步鉴定了结缕草属植物5种1变种97份种源叶片抗寒性,在此基础上,选取其中15份抗寒性有差异的种源的匍匐茎和根状茎进行生长恢复试验。结果表明：(1)以叶片半致死温度(LT50)为评价指标,结缕草属植物的LT50的范围是-1.9℃～-10.4℃,变异系数为27.79%；结缕草属植物不同种的LT50从低到高依次为：结缕草<中华结缕草<沟叶结缕草(Z.matrella (L.) Merr.)<细叶结缕草(Ztenuifolia Thiele.)<长花结缕草(Z.sinica var longiflora Ohwi.)<大穗结缕草(Zmacrostachya Franchet et Sav.);(2)结缕草、中华结缕草和沟叶结缕草三个种种内LT50的变异都很大,变异系数(CV)都高于20%。(3)不同季节的匍匐茎和根状茎的LT50间存在显著的正相关。(4)不同器官的LT50间的R均达到了显著水平。
4.2结缕草冷驯化条件下可溶性蛋白变化分析
通过对极端抗寒性的两个结缕草种源Z125(弱)和Z141(强)的叶片、匍匐茎和根状茎三个器官在正常生长条件(28℃/22℃,白天/黑夜,8h光照)与模拟低温驯化(8℃/2℃,白天/黑夜,光照8h)后的可溶性蛋白的SDS-PAGE电泳分析,发现,在模拟低温处理后2周,抗寒性差的Z125的可溶性蛋白合成发生较大的变化,根状茎比对照多了44KD的一条带,叶片比对照多了66KD左右的带,而抗寒性强的Z141仅在根状茎中比对照多了一条50KD的带,但少了一条40KD左右的带；处理4周后,Z141的匍匐茎和根状茎的可溶性蛋白都比对照多了两条带,大小为60KD左右,其叶片的可溶性蛋白电泳条带比未驯化的少了2条带(分别是60KD和20KD左右),而Z125仅根状茎可溶性蛋白比对照多了两条带,其多肽大小约为50KD。
5结缕草属(Zoysia Willd.)植物遗传变异和种间关系
5.1形态学研究
通过对结缕草属植物9种1变种59份种源的23个外部性状的观测和分析,得出结论：①沟叶结缕种内各种源间在很多性状上都存在很大的变异,结缕草和中华结缕草的种内变异也较大,次于沟叶结缕草,并且两者的种内变异大小较接近。朝鲜结缕草(Z. koreana Mez)种内变异小；②这些种之间根据遗传距离远近可以分成三大类：第一类为结缕草、中华结缕草、朝鲜结缕草先聚在一起之后再和大穗结缕草聚成一大类；第二类为沟叶结缕草和细叶结缕草先聚为一小类,长花结缕草和小结缕草(Zminima Zotov)聚为另一小类,然后这些种聚成一大类。大花结缕草(Z. macrantha Desvaux)独自成一集群为第三类；③主成分分析结果表明：叶片宽度,匍匐茎长度、直径,每穗小穗数,小穗柄长度,小穗长度,生殖枝高度,叶舌的颜色及有无毛这9个性状对结缕草属植物种间亲缘关系的研究具有重要的意义。
5.2等位酶研究
分析结缕草属植物9种1变种59份种源的苹果酸脱氢酶(MDH),酯酶(EST)及过氧化物酶(POD)三种等位酶特征,研究其种间亲缘关系,结果表明：①种内遗传分化大小依次为：沟叶结缕草>结缕草>中华结缕草>朝鲜结缕草；②聚类结果显示：结缕草、中华结缕草、长花结缕草、大穗结缕草聚为第一类；沟叶结缕草、细叶结缕草先聚为一类,然后小结缕草、太平洋结缕草(Z. pacifica Hotta)、大花结缕草也聚到这一类；而朝鲜结缕草自成一类。
5.3SSR分子标记研究
利用SSR分子标记对结缕草属植物9种1变种59份种源进行了种间关系分析,结果表明：①种内遗传分化大小依次为：结缕草、中华结缕草>朝鲜结缕草>沟叶结缕草；②聚类结果显示：结缕草、中华结缕草、朝鲜结缕草和大穗结缕草聚为一类；沟叶结缕草、小结缕草和细叶结缕草三个种聚为一类；大花结缕草、长花结缕草各自为一类。
6中国结缕草属种质资源综合评价
应用叶色、密度、叶宽、均一性、青绿期、抗寒性、锈病抗性7个指标,对中国结缕草属植物的74份种源在南京地区的坪用价值进行评价,结果表明：Z014、Z075、Z008、Z142、Z042、Z005、Z044、Z101、Z053、Z050等种源叶色较深而密度较高,青绿期较长但抗寒性一般,综合坪用质量高；而Z118、Z081、Z010、Z115、Z104、Z103等种源叶色黄绿,青绿期较短,抗寒性较差,锈病抗性差,综合坪用质量低。其他54份种源由于2个或3个指标得分较低,综合坪用质量中等。
Zoysia grasses (Zoysia spp.) are widely used as warm season turf grass indigenous to the countries of the western Pacific Rim. They possess excellent wear tolerance, drought tolerance, disease resistance, salinity tolerance, low fertility tolerance and low maintenance requirements.
Forteen morphological characteristics, phonological phase, seed yield and its component, stolon growth rate of representative accessions of5species and1variety of Zoysia Willd. were observed in this report. The leaf low temperature of97accessions of5species and variety of Zoysia Willd. was evaluated using modified electrolyte leakage method, and15representative accessions were selected based on leaf low temperature result to evaluate the low-temperature tolerance of stolons and rhizomes using survival rate, and then soluble protein of three organs of two extreme low temperature tolerant representative accessions were studied using SDS-PAGE electrophoresis. Morphology, isozyme and SSR genetic markers at three different levels were studied combining with the geographical distribution to study the genetic variation and interspecific relationship of59accessions of Zoysia Willd. belonging to9species and1variety. The study will provide an important scientific basis for the development and untilization of zoysiagrass germplasm resource in China.
The main results were stated as followings:
1. Variation analysis on morphological characteristic of Zoysia Willd. in China
The variations and tendencies of14morphological and their correlationship of124accessions of Zoysia Willd. were studied in the paper. The results showed that the great variation of morphological characteristics were existed among the accessions of Zoysia Willd.with the coefficient of variation (CV) at exceed10%. Correlation analysis indicated obvious correlation were existed among some of14morphological characteristics at significant levels. Some morphological characteristics was correlated siginificantly with latitude or longitude. Four groups were divided based on14morphological characteristics: fine zoysiagrass, inflorescenced coarse zoysiagrass, less-inflorescenece coarse zoysiagrass and Z099.
2. Study on the variation of phenological phase of Zoysia Willd.
The phenological phases of109accessions of Zoysia Willd. including5species and1variety were observed. The results showed that there is difference in the date of phonological phases among different accessions in Nanjing. Variation scopes of blooming stage and wilting stage was biggest, and that of green period was215-277d. The date of initial booting stage has a significantly positive correlation with the date of initial blooming stage, blooming stage, maturation stage and wilting stage and length of green period. The date of wilting stage has a positive correlation with the length of green period, and that of turning-green stage has a highly significant negative correlaion with the length of green period. Regression analysis indicated that turing green, initial booting, initial blooming, blooming and wilting stages of Zoysia Willd. became obviously earlier with latitude increasing. The phonological phase of coarse texture accessions was earier than fine texture accessions relatively.
3. Study on propagation properties of Zoysia Willd. in China
The stolon growth rate, the seed yield and its component of Z. japonica and Z. sinica in China were measured and analyzed. The results showed that in Nanjing:(1) abundant variation was existed in the total stolon length and aboveground biomass of different accessions of zoysiagrass;(2) the total stolon length was positive corrected with the aboveground biomass with correlation coefficient as0.676;(3) abundant variation was existed in the seed yield and its component, in which seed yield had the highest variation with the coefficient vatiation (CV) as102.51%;(4) seed number per spike and the weight per100seeds were positive corrected with seed yield, inflorescence density was not corrected with seed yield directly, but seed yield was correlated with inflorescence density, seed number per spike and the weight per100seeds after they interacted;(5) path analysis showed that seed number per spike and the weight per100seeds contributed the most to the seed yield, inflorescence density contributed few;(6) above ground biomass was negatively correlated siginificantly with inflorescence density;(7) the multiple comparisons analysis of the total stolon length and seed yield revealed the vegetative growth rate of six accession such as Z015,Z013, Z056et al was high, nine accessions such as Z016, Z052and Z061et al have high seed yield, six accessions such as Z012, Z063et al have high seed yield and vegetative growth rate, and the vegetative growth and seed yield of five accessions such Z111, Z018et al was both low.
4. Study on Low Temperature Tolerance of Zoysia Willd.
4.1The assessment of the Low Temperature Tolerance of Zoysia Willd.
The leaf LT50of97accessions of5species and1variety of Zoysia Willd. were evaluated by electrolyte leakage, then stolon and rhizome low temperature of15representative accession were evaluated by survival rate. The leaf LT50of Zoysia Willd was from-1.9℃to-10.4℃with coefficient of variation (CV) of27.79%, and the LT50of different species of Zoysia Willd were ranked as follows:Z.japonica.
Soluble proteins in leaves, stolon and rhizome of Z125(poor cold tolerance) and Z141(strong cold tolerance) in non-acclimation (28℃/22℃, day/night,8h light) and simulated low temperature acclimation(8℃/2℃, day/night,8h light) were studied. SDS-PAGE electrophoresis revealed that after two weeks of cold acclimation, the soluble protein synthesis in Z125had a major change, a new40KD protein components in rhizome and a66KD in leaves in cold-acclimation comparing to the non-acclimation; a more50KD and less40KD protein components changed in rhizome in cold-acclimation Z141comparing to non-acclimation Z141.After4weeks of cold-acclimation, two new protein compositions, about60KD, appeared in cold-acclimated Z141both stolon and rhizome, about60KD and20KD soluble protein compositions were disappeared in cold-acclimated Z141leaf. And only2new soluble protein compositions,about50KD, appeared in cold-acclimated Z125rhizome.
5. Study on Genetic Variation and Interspecific Genetic Relationship of Zoysia Willd.
5.1The morphological markers
With observing and measuring23morphological characteristics on59accessions of Zoysia Willd., the results showed as:The accessions of Z.matrella had greatest variation on most characters, Z.japonica and Z.sinica had great intraspecies differentiation but smaller than Z.matrella, and the degree of genetic diversity within species of these two were very close. The accessions of Z.koreana Mez. had little differentiation on many characters respectively, and the genetic diversity of Z.koreana were smaller;②The cluster analyzing results show:All these species can be classified by genetic distance to be3classes:the first class included of Z. japonica and Z sinica, Z. koreana and Z. macrostachy., the second class included Z. tenuifolia, Z. matrella, Z. minima Zotov. and Z.sinica var. longiflora, Z. macrantha was clustered to be a group itself;③The principal components analyzing results showed that the width of blades, the length and diameter of stolons, the spikelets number of one spike, the length of spikelets and the pedicels, the height of procreant culms, the ligule colour, whether ligule with hairs or not, these features are important in researching of the genetic relationship of species.
5.2Isozyme study
Three isozyme systems (MDH、EST、POD) were conducted on59accessions of Zoysia Willd. to study the genetic relationship of species. The results show as follows:①the genetic diversity was ranked as:Z.matrella> Z. japonica> Z.sinica> Z.koreana.;②the clustering analysis results show:Z. japonica and Z. sinica> Z. sinica var. longiflora and Z.macrostachya were clustered together; Z.matrella、Z. tenuifolia were clustered to be a group, then Z.minima、Z. pacifica、Z. macrantha were also clustered to this group; Z.koreana was clustered to be a group by itself.
5.3SSR analysis
SSR markers were used to study the genetic relationship of accessions of Zoysia Willd..①The genetic diversity within species ranked as:Z.japonica, Z. sinica> Z. koreana> Z. matrella;②Z.japonica was clustered together with Z. sinica, Z. macros tachya and Z koreana; Z. matrella was clustered together with Z. tenuifolia and Z. minima; Z. macrantha and Z.sinica var. longiflora was a group by itself respectively.
6. The evaluation of turf value of Zoysia Willd. germplasm in China
The turf value of74accession of Zoysia Willd.in China were evaluated based on leaf color, density, leaf width, uniformity, green period, low temperature tolerance and rust resistance index. The result showed that ten accessions including Z014、Z075、Z008、Z142、 Z042、Z005、Z044、Z101、Z053and Z050were dark-green, dense, long green-period and common low temperanture tolerance, and the turf quality of these accessions were very high. Ten accessions including Z118、Z081、Z010、Z115、Z104、and Z103et al were yellow-green, short green-period and low tolerancc to low temperature and rust, the turf quality of these ten accessions was very low. The the turf value of other54accessions were intermediated due to two or three poor index.
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