Construction of a barley (Hordeum vulgare L.) YAC library and isolation of a Hor1-specific clone

We have constructed an EcoRI-based YAC (yeast artificial chromosome) library from barley (Hordeum vulgare L. cv. Franka) using the vector pYAC4. The library consists of approximately 18 000 recombinant YACs with insert sizes ranging between 100 and 1000 kb (average of 160 kb) corresponding to 50% of the barley genome. Size fractionation after ligation resulted in an increased average insert size (av. 370 kb) but also in a substantial decrease in cloning efficiency. Less than 1% of the colonies showed homology to a plastome-specific probe; approximately 50% of the colonies displayed a signal with a dispersed, highly repetitive barley-specific probe. Using a primer combination deduced from the sequence of a member of the small Hor1 gene family coding for the C-hordein storage proteins, the library was screened by polymerase chain reaction and subsequently by the colony hybridization technique. A single YAC, designated Y66C11, with a 120 kb insert was isolated. This DNA fragment represents a coherent stretch from the terminal part of the Hor1 gene region as judged from the correspondence of the restriction patterns between Y66C11 DNA and barley DNA after hybridization with the Hor1-specific probe. Restriction with the isoschizomeric enzymes HpaII/MspI suggests a high degree of methylation of the Hor1 region in mesophyll cells but not in YAC-derived (yeast) DNA.     

青稞HvBAS1基因的克隆及表达分析

细胞色素P450单加氧酶(CYP450)是参与植物代谢的最大酶家族,其中CYP734A亚家族成员广泛参与植物激素油菜素类固醇(BRs)的失活。该研究以青稞农家品种‘肚里黄’幼苗为实验材料,通过人工合成激素24 表油菜素内酯(24 eBL)和BRs合成抑制剂油菜素唑(BRZ)处理,分析BRs对青藏高原特色作物青稞(Hordeum vulgare L. var. nudum Hook. f.)的影响;采用RT PCR技术从青稞中克隆HvBAS1基因,并运用实时定量PCR检测其表达特征,为深入分析HvBAS1基因的功能奠定研究基础。结果显示：(1)24 eBL能够显著促进青稞幼苗的生长,而BRZ处理后幼苗长势明显减缓。(2)从青稞中成功克隆到2个与拟南芥BRs失活基因BAS1高度同源的CYP734A亚家族基因,即HvBAS1 1和HvBAS1 2;HvBAS1 1开放阅读框全长1 629 bp,编码542个氨基酸;HvBAS1 2长1 689 bp,编码562个氨基酸,二者氨基酸序列相似性为76.42%;亚细胞定位预测结果显示二者均存在于内质网。(3)实时定量PCR检测发现,青稞HvBAS1 1与HvBAS1 2的表达模式完全不同,其中HvBAS1 1在青稞根中的表达量高于叶中,而HvBAS1 2在叶中的表达量高于根,且随着苗龄的增大,HvBAS1 1在根中的表达呈先升高后降低的趋势,HvBAS1 2在叶片中呈先降低后升高的变化趋势;BRZ处理青稞幼苗后,其HvBAS1 1与HvBAS1 2基因的表达较对照均显著下调,且HvBAS1 2下调更为明显。研究表明,HvBAS1 1和HvBAS1 2很可能都参与了青稞内源BRs的失活,但二者的功能存在差异。     

The inheritance of volatile phenylpropenes in bitter fennel (Foeniculum vulgare Mill. var. vulgare, Apiaceae) chemotypes and their distribution within the plant

The volatile phenylpropenes estragole and t-anethole are the major constituents of the oleoresin of the aerial parts of bitter fennel (Foeniculum vulgare Mill. var. vulgare, Apiaceae). The levels of estragole and t-anethole varied during plant development, being maximal in flowers and developing mericarps. Still the ratio between estragole and t-anethole remained constant throughout development. Estragole-rich types were hybridized with t-anethole rich types to examine the genetic basis of this polymorphism. A reverse correlation between estragole and t-anethole content was evident and the action of a biallelic gene with partial dominance for high estragole content was inferred. Understanding phenylpropene inheritance might explain chemical polymorphism in wild bitter fennel populations, sheds light on the molecular mechanisms that lead to chemotypes evolution and is crucial for breeding fennel varieties with desired chemical compositions.     

黄果龙葵和龙葵染色体制片优化及核型分析

该实验以黄果龙葵和龙葵的根尖为实验材料,进行不同的预处理、固定和解离,确定出各种材料适合于核型分析的制片方法。结果表明:龙葵于15℃条件下经0.05%秋水仙素预处理2.5h,固定后用1mol/L HCl酸解后,染色观察,得到的染色体分散,易于染色体计数和形态观察。用此方法对黄果龙葵和龙葵进行核型分析,结果发现:黄果龙葵和龙葵都属于小型染色体,黄果龙葵为四倍体,核型公式为K(4n)=48=4sm+44m,核型不对称系数为56.22%,属于2B核型。龙葵为六倍体,核型公式为K(6n)=72=72m,核型不对称系数为55.89%,属于1B核型。     

利用粳稻染色体片段置换系群体检测水稻抗亚铁毒胁迫有关性状QTL

潜育性水稻田广泛分布于中国、斯里兰卡、印度、印度尼西亚、塞拉里昂、利比亚、尼日利亚、哥伦比亚和菲律宾等国,其中我国南方稻区就有近700万公顷低产潜育性水稻田。该类水稻田还原性强,矿质营养失调,尤以Fe^2 过量积累,对水稻生长发育产生不良的逆境胁迫作用。培育抗亚铁毒的水稻品种是简便、经济有效地提高稻谷产量的重要途径之一。该文利用由粳稻品种Asominori与籼稻品种IR24杂交衍生的Asominori染色体片段置换系(Chromosome Segment Substitution Lines,CSSLs)群体为材料,检测与抗亚铁毒胁迫有关性状QTL。共检测到与抗亚铁毒胁迫有关性状QTL14个,各QTL的LOD值为2．72～6．63。其中检测到与抗亚铁毒胁迫直接有关的性状叶片棕色斑点指数QTL3个,分别位于第3、9、11染色体C515～XNpb279、R2638～C1263和G1465～C950之间,对应的贡献率分别为16．45％、11．16％和28．02％;与其他已发表的定位结果比较发现,位于第三染色体C515～XNpb279间控制叶片棕色斑点指数的QTL与水稻功能图谱上控制叶绿素含量的QTL的位置一致;表明在亚铁毒胁迫条件下,水稻在其叶片表面出现棕色斑点,叶片衰老,产生一些叶绿素降解物或衍生物,以提高叶片细胞对亚铁等重金属毒害的耐受力。另外,在第11染色体G1465～C950之间检测到了控制叶片棕色斑点指数、茎干重和根干重QTL1个,为主效QTL。在第6染色体XNpb386～XNpb342之间检测到控制茎干重、株高、根长和根干重QTL1个,是否与水稻抗亚铁毒有关需要进一步研究。本研究旨在通过定位与抗亚铁毒有关的QTL,借助与之紧密连锁的分子标记有效地聚合这些QTL,培育出抗亚铁毒性强的水稻新种质材料。     

Expression of sucrose: fructan 6-fructosyltransferase (6-SFT) and myo -inositol 1-phosphate synthase (MIPS) genes in barley (Hordeum vulgare) leaves

Fructan is an important class of non-structural carbohydrates present in cool-season grasses. Sucrose: fructan 6-fructosyltransferase (6-SFT, EC 2.4.1.10), one of the enzymes thought to be involved in grass fructan biosynthesis, catalyzes the initiation and extension of 2,6-linked fructans.Myo-inositol is a central component in several metabolic pathways in higher plants.Myo-inositol 1-phosphate synthase (MIPS) (EC 5.5.1.4), the first enzyme in inositolde novo biosynthesis, catalyzes the formation ofmyo-inositol 1-phosphate (MIP) from glucose-6-phosphate. The expression of 6-SFT and MIPS genes is compared in barley (Hordeum vulgare L.) leaves under various conditions. In cool temperature treatments, both 6-SFT and MIPS mRNAs accumulate within two days and then decline after four days. Under warm temperatures and continuous illumination, the amount of 6-SFT and MIPS mRNA gradually accumulated in detached leaves and increased significantly by 8 h. In contrast, we observed no significant changes over time in attached (control) leaves. Treating detached leaves with glucose or sucrose in the dark resulted in accumulations of both 6-SFT and MIPS mRNA. Homologous expression patterns for 6-SFT and MIPS genes suggest that they may be similarly regulated in barley leaves. Although sucrose and glucose may play important roles in the expression of 6-SFT and MIPS genes, regulation likely involves multiple factors.     

2′-hydroxymethyl abscisic acid: A major catabolite of (±)-abscisic acid in leaves of Hordeum vulgare L.

Radioactive (±)-abscisic acid (ABA), supplied via the transpiration stream to light-grown leaves of Hordeum vulgare was catabolized to 2′-hydroxymethyl ABA. Identification was made by capillary gas chromatography-mass spectrometry (GC-MS).     

QTL Analysis of Aluminum Resistance in Rice (Oryza sativa L.)

Aluminum (Al) toxicity is considered as one of the primary causes of low-rice productivity in acid soils. In the present study, quantitative trait loci (QTLs) controlling Al resistance based on relative root elongation (RRE) were dissected using a complete linkage map and a recombinant inbred lines (RILs) derived from a cross of Al-tolerant japonica cultivar Asominori (Oryza sativa L.) and Al-sensitive indica cultivar IR24 (O. sativa L.). A total of three QTLs (qRRE-1, qRRE-9, and qRRE-11) were detected on chromosomes 1, 9, and 11 with LOD score ranging from 2.64 to 3.60 and the phenotypic variance explained from 13.5 to 17.7%. The Asominori alleles were all associated with Al resistance at all the three QTLs. The existence of these QTLs was confirmed using Asominori chromosome segment substitution lines (CSSLs) in IR24 genetic background (IAS). By QTL comparative analysis, the two QTLs (qRRE-1and qRRE-9) on chromosomes 1 and 9 appeared to be consistent among different rice populations while qRRE-11 was newly detected and syntenic with a major Al resistance gene on chromosome 10 of maize. This region may provide an important case for isolating genes responsible for different mechanisms of Al resistance among different cereals. These results also provide the possibilities of enhancing Al resistance in rice breeding program by marker-assisted selection (MAS) and pyramiding QTLs.     

The Nir1 locus in barley is tightly linked to the nitrite reductase apoprotein gene Nii

pBNiR1, a cDNA clone encoding part of the barley nitrite reductase apoprotein, was isolated from a barley (cv. Maris Mink) leaf cDNA library using the 1.85 kb insert of the maize nitrite reductase cDNA clone pCIB808 as a heterologous probe. The cDNA insert of pBNiR1 is 503 by in length. The nucleotide coding sequence could be aligned with the 3 end of other higher plant nitrite reductase apoprotein cDNA sequences but diverges in the 3 untranslated region. The whole-plant barley mutant STA3999, previously isolated from the cultivar Tweed, accumulates nitrite after nitrate treatment in the light, has very much lowered levels of nitrite reductase activity and lacks detectable nitrite reductase cross-reacting material due to a recessive mutation in a single nuclear gene which we have designated Nir1. STA3999 has the characteristics expected of a nitrite reductase apoprotein gene mutant. Here we have used pB-NiR1 in RFLP analysis to determine whether the mutation carried by STA3999 is linked to the nitrite reductase apoprotein gene locus Nii. An RFLP was identified between the wild-type barley cultivars Tweed (major hybridising band of 11.5 kb) and Golden Promise (major hybridising band of 7.5 kb) when DraI-digested DNA was probed with the insert from the partial barley nitrite reductase cDNA clone, pBNiR1. DraI-digested DNA from the mutant STA3999 also exhibited a major hybridising band of 11.5 kb after hybridisation with the insert from pBNiR1. F₁ progeny derived from the cross between the cultivar Golden Promise and the homozygous nir1 mutant STA3999 were heterozygous for these bands as anticipated. Co-segregation of the Tweed RFLP band of 11.5 kb and the mutant phenotype (leaf nitrite accumulation after nitrate treatment/loss of detectable nitrite reductase cross-reacting material at M_r 63000) was scored in an F₂ population of 312 plants derived from the cross between the cultivar Golden Promise and the homozygous mutant STA3999. The Tweed RFLP band of 11.5 kb and the mutant phenotype showed strict co-segregation (in approximately one quarter (84) of the 312 F₂ plants examined). Only those F₂ individuals heterozygous for the RFLP pattern gave rise to F₃ progeny which segregated for the mutant phenotype. We conclude that the nir1locus and the nitrite reductase apoprotein gene Nii are very tightly linked.     

Plant development and synthesis of essential oils in micropropagated and mycorrhiza inoculated plants of Origanum vulgare L. ssp. hirtum (Link) Ietswaart

Biomass production of micropropagated oregano was induced by inoculation with the fungus Glomus viscosum. The effects of arbuscular mycorrhizal (AM) symbiosis on morphological and metabolic variations of regenerated oregano plants were investigated at different growth stages. AM greatly increased parameters such as plant leaf area, fresh and dry weight, number of spicasters and verticillasters in infected plants. An increase of the gland density, especially on the upper leaf epidermis, was also observed following the physiological ageing of the tissues. The in vitro plants of O. vulgare ssp. hirtum described in this study provided a qualitatively and quantitatively good source of essential oils that have a chemical profile comparable to that of the control mother plants with carvacrol as the main compound.     

不同产地木耳菜的染色体核型分析

为了研究木耳菜核型特征及不同产地间的进化关系,以来自7个产地的8个木耳菜品种为材料,采用常规压片法进行核型分析,并进行核型进化趋势分析和主成分分析。结果表明:(1)所有木耳菜的染色体数目均为2n=2x=44,未见异常染色体,染色体类型均为中部着丝粒染色体(m)或近中部着丝粒染色体(sm),且m数量多于sm。(2)不同产地的木耳菜在染色体核型公式、核型类型、随体位置、染色体长度比、臂比及核型不对称系数等指标均存在明显差异;随体均为1对,但随体位置不同。(3)核型类型为1A、1B和2A型,其中1A型5种,数量最多。(4)染色体长度比范围为1.51~2.06,平均臂比值范围为1.30~1.48,仅有吉林‘利丰’和江西‘航城’存在臂比大于2的染色体。(5)核型不对称系数范围为56.25%~59.17%,核型的对称程度较高,推测木耳菜的进化程度较为原始,其中河北‘金发’是最原始,江西‘航城’最进化。研究结果为木耳菜的细胞遗传学研究提供了参考依据。     

一种由粉红粘帚霉引起的青稞根腐病

[背景]根腐病在青稞生产中的危害日趋严重,阻碍了青稞根腐病的有效防控及青海省青稞产业的发展。然而人们对青稞根腐病的研究甚少且病原菌不详。[目的]明确青稞根腐病发生的危害、病原及致病性,为青稞根腐病的防控提供理论依据。[方法]采用常规的组织分离法分离青稞根腐病病原,通过形态鉴定与分子鉴定结合的方法对病原进行鉴定,并采用烧杯水琼脂法测定其致病性。[结果]共分离得到4株青稞根腐病病原菌,鉴定为Clonostachys rosea,有较强的致病性且致病性差异显著,经柯赫氏法则验证为青稞根腐病病原菌,并且是一种新的青稞根腐病病原,该类根腐病也是一种新的根腐类病害,在国内外属首次发现。[结论]Clonostachys rosea可引起青稞根腐病且致病性强。     

Flt-2L, a locus in barley controlling flowering time, spike density, and plant height

Flowering time represents an important adaptive trait for temperate cereal crops and may also impact on frost damage in cereal reproductive tissues by enabling escape or by influencing accumulation of genuine tolerance. The Flowering time-2L (Flt-2L) quantitative trait locus (QTL) on the distal end of barley chromosome arm 2HL overlaps with QTL for rachis internode length and reproductive frost damage. Flt-2L was also found to be associated with plant height. By combining marker analysis with phenotyping in progeny families of selected Amagi Nijo × WI2585 F₆ recombinants, we were able to map quantitative flowering time, rachis internode length, and plant height effects on 2HL as discrete Mendelian traits. The three developmental characters showed codominant modes of expression and perfectly cosegregated with one another in a 1.3-cM marker interval, indicating control by the same gene or closely linked genes. Twelve genes were identified in the related intervals in the rice and Brachypodium distachyon genomes. The HvAP2 gene cosegregated with Flt-2L and represents a plausible candidate for Flt-2L, since it is highly similar to the wheat domestication gene Q which has similar developmental effects. These data will contribute to isolation of the Flt-2L gene(s) and help establish the basis of the frost damage QTL. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

小桐子糖原合成激酶基因JcGSK的克隆与表达分析

以小桐子(Jatropha curcas L.)cDNA为模版,克隆了JcGSK基因的CDS序列。序列分析表明,JcGSK基因包含1 230bp完全阅读框(ORF),编码409个氨基酸。预测其编码蛋白质的相对分子量为46.33kD,理论等电点为8.58。Blast搜索结果及进化分析结果表明,JcGSK蛋白与巴西橡胶树GSK蛋白的氨基酸序列一致性最高(94%)且亲缘关系最近;JcGSK基因编码的蛋白具有一个蛋白激酶特有的结构域。组织表达结果显示,JcGSK基因在小桐子根、茎、叶、花、果皮和种子中都有表达,且在根中表达量最高。小桐子幼苗在NaCl、ABA、PEG、低温和机械损伤处理后JcGSK基因表达量有不同程度的上调,推测其参与小桐子非生物胁迫响应和信号传导过程。JcGSK基因在种子中也有较高表达,在种子发育过程中表达量的变化与种子生长发育趋势基本一致,推测JcGSK基因也参与调控小桐子种子的生长发育。     

沙鞭不同居群染色体数目及核型分析

该研究采用常规压片法对分布于内蒙古高原的6个沙鞭居群的染色体核型进行研究,探讨沙鞭不同居群的核型特征及其进化关系。结果表明：(1)沙鞭6个居群的染色体数目恒定,均为2n = 2x = 46。(2)染色体有正中部着丝粒(M)、中部着丝粒(m)、亚中部着丝粒(sm)和亚端部着丝粒(st)4种类型,且中部着丝粒类型数量最多。(3)沙鞭不同居群核型公式存在差异。(4)核型类型有1A、2A、1B和2B型4种,染色体平均臂比介于1.29 ~ 1.62,长度比为1.73 ~ 2.68。(5)核型不对称系数处于55.96% ~ 59.95%,核型对称性较高,进化程度较为原始,其中37居群的核型不对称系数最大,进化程度较高,34居群的核型不对称系数最小,进化程度较低。(6)沙鞭6个居群聚为两类,37居群单独聚为一类,其他所有居群聚为一类,表明37居群与其他居群具有相对较远的亲缘关系。该研究首次报道了沙鞭不同居群的染色体核型特征及其进化关系,为以后沙鞭的系统进化和优良种质资源筛选奠定了细胞学证据。     

石榴PgUFGT基因克隆及表达分析

利用RT-PCR和RACE方法,从石榴(Punica granatum L.)果皮中克隆到一个类黄酮糖基转移酶(UFGT)基因(PgUFGT)全长cDNA序列(GenBank登录号为KF841620)。PgUFGT基因编码区1 476bp,编码491个氨基酸。PgUFGT蛋白具有保守PSPG基序、UDP-糖基转移酶家族结构域和UDP-葡萄糖醛酸基/葡萄糖基转移酶保守域(UDPGT),与其他植物UFGT蛋白一致性较高;系统进化树分析结果表明,PgUFGT属于类黄酮3-O-糖基转移酶类。荧光定量qRT-PCR结果表明,PgUFGT基因在‘红宝石’和‘水晶甜’2个石榴品种的发育期内具有不同的表达模式,PgUFGT在‘红宝石’石榴中有2个转录表达高峰,而在‘水晶甜’石榴中仅有1个表达高峰,表明PgUFGT可能在2个石榴品种中具有不同的催化作用。该研究结果为进一步研究石榴果实色泽形成的分子机制奠定了基础。     

白菜型油菜RbohC和RbohF基因克隆与表达分析

该研究以白菜型油菜（Brassica rapa L.）‘陇油6号’为实验材料,采用RT PCR方法克隆油菜RbohC和RbohF基因,并采用实时荧光定量PCR技术对RbohC、RbohF基因在不同组织及非生物胁迫下的表达进行分析,为深入研究油菜RbohC、RbohF基因的生物学功能提供依据。结果显示：(1) 成功克隆得到2个全长分别为3 050 bp和2 995 bp的油菜RbohC (GenBank登录号：XM_009134386) 和RbohF (GenBank登录号：XM_009114548) 基因序列。(2) 生物信息学分析显示,油菜RbohC和RbohF基因开放阅读框（ORF）分别为2 733 bp和2 847 bp,编码910和948个氨基酸,推测二者的蛋白质分子量分别为103 kDa和108 kDa,理论等电点分别为9.47和9.21; 油菜RbohC、RbohF编码的氨基酸序列与萝卜等多种植物相应蛋白氨基酸序列具有较高的同源性,且这些序列高度保守并含有NADPH氧化酶的典型保守结构域,包括2个可以与Ca²⁺结合的EF手性模体结构、6个跨膜结构域、黄素腺嘌呤二核苷酸结合结构域、NAD焦磷酸结合结构域和C末端区域中的NADP核糖保守结合位点。(3) 油菜RbohC和RbohF基因在根、茎、叶和下胚轴中均表达,无组织特异性,但RbohC基因在根中表达量最高, RbohF基因在下胚轴中表达量最高。(4) 低温、干旱、盐、ABA、H₂O₂处理都能够诱导油菜RbohC和RbohF基因的表达,但抗寒性强的 ‘陇油6号’的RbohC和RbohF基因对胁迫的响应更敏感,且RbohC基因的表达量均高于RbohF基因。(5) 用H₂O₂清除剂DMTU、NADPH氧化酶抑制剂DPI和IMD、MAPKK抑制剂U0126处理后,油菜RbohC和RbohF基因的表达均较对照下降,说明U0126和DMTU对油菜RbohC和RbohF基因的表达有抑制作用。研究认为,油菜RbohC和RbohF基因在油菜适应逆境胁迫中具有重要作用,两基因的表达均受MAPK激酶信号途径的调节,并受到H₂O₂的反馈调节,而且抗寒性强的‘陇油6号’品种中RbohC和RbohF基因对H2O2和MAPK激酶信号途径的响应更敏感。     

Cloning and Characterization of Four B-hordein Genes from Tibetan Hull-less Barley (Hordeum vulgare subsp. vulgare)

Four B-hordein genes, designated BH1-BH4, were cloned using PCR amplification from two hull-less barley cultivars, ZQ7239 and ZQ148, collected from Tibet. The results of sequencing indicated that BH1-BH4 contained complete open reading frames (ORFs). Comparison of their predicted polypeptide sequences with the published sequences suggested that they all share the same basic protein structure. Phylogenetic analysis indicated that the deduced amino-acid sequences of BH1-BH4 genes were more closely related to B-hordeins from cultivated barley (Hordeum vulgare L.) than to any other prolamins from wild barley and Aegilops tauschii. Comparison of the coding regions of BH1-BH4 genes showed that BH1 had a lower sequence identity to other previously published B-hordeins than the other three B-hordeins obtained in this study. BH1 was then cloned in a bacterial expression vector based on bacteriophage T7 RNA polymerase. The resulting plasmid produced a 28.15 kDa protein in Escherichia coli. The potential value of B-hordein genes in grain quality improvement of hull-less barley has been discussed.     

呼伦贝尔黄花苜蓿MfMYB30基因的克隆及表达分析

该研究在呼伦贝尔黄花苜蓿(Medicago falcata L. cv. Hulunbuir)转录组测序基础上,通过RT PCR方法克隆获得了MfMYB30基因,并通过生物信息学和表达分析进行初步研究,为深入研究MfMYB30基因的功能和开发利用奠定基础。结果表明：(1)成功克隆获得呼伦贝尔黄花苜蓿MfMYB30基因,其ORF序列长为957 bp,编码318个氨基酸,相对分子质量为86.85 kD,理论等电点为5.11。MfMYB30蛋白为疏水性蛋白,无跨膜结构,无信号肽序列。(2)系统进化分析表明黄花苜蓿MfMYB30蛋白与紫花苜蓿MsMYB4、拟南芥AtMYB30、木豆CcMYB30、蒺藜苜蓿MtMYB30和大豆GmMYB60聚为一个类群,亲缘关系较近。(3)实时荧光定量PCR结果显示,MfMYB30在黄花苜蓿模拟刈割不同天数后的相对表达量呈先降低后升高的趋势,在刈割7 d后相对表达量达到峰值。(4)通过在拟南芥原生质体亚细胞定位分析发现,该蛋白定位于细胞核。研究推测,MfMYB30基因可能在黄花苜蓿刈割或放牧胁迫响应过程中发挥重要调控作用。