Genetic characteristics and research advances of genes related to cotton fiber development

MAO Wei; CAO Yuefen

doi:10.11833/j.issn.2095-0756.2018.06.021

Volume 35 Issue 6

Nov. 2018

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MAO Wei, CAO Yuefen. Genetic characteristics and research advances of genes related to cotton fiber development[J]. Journal of Zhejiang A&F University, 2018, 35(6): 1155-1165. doi: 10.11833/j.issn.2095-0756.2018.06.021

Citation:

MAO Wei, CAO Yuefen. Genetic characteristics and research advances of genes related to cotton fiber development[J]. Journal of Zhejiang A&F University, 2018, 35(6): 1155-1165. doi: 10.11833/j.issn.2095-0756.2018.06.021

Genetic characteristics and research advances of genes related to cotton fiber development

doi: 10.11833/j.issn.2095-0756.2018.06.021

MAO Wei^1
,,
CAO Yuefen^{1,2
,
,}

1.
The Key Laboratory for Quality Improvement of Agricultural Products of Zhejiang Province, School of Agriculture and Food Science, Zhejiang A & F University, Hangzhou 311300, Zhejiang, China
2.
Zhejiang Key Laboratory of Crop Germplasm, College of Agriculture and Biotechnology, Zhejiang University, Hangzhou 310058, Zhejiang, China

Received Date: 2017-11-15
Rev Recd Date: 2018-01-28
Publish Date: 2018-12-20

Abstract

Cotton fiber is a major raw textile material which plays an essential role in the national economic development. The developmental process of cotton fiber cell is an orderly and systematically multi-gene controlled process including the fiber initiation, elongation, secondary wall thickening and the finally maturation process. With highly integrated studies of genetics, cytology and molecular biology, the molecular mechanisms of cotton fiber growth have become a popular focus both abroad and domestically. Most of the research is on the genetic mapping, gene cloning and high throughout sequencing areas in recent years. In order to systematically understand the development of cotton fibers, this paper summarized the morphological changes and characteristics during cotton fiber development and the classical genetics mapping research on the cotton fiber related genes, and took an overview of the application and progress of deep sequencing technology in cotton fiber development from transcriptome, proteomics and epigenetics field, and untangled the related genes in different periods of cotton fiber development. The previous studies had proved that the fiber yield was determined by the initial stage of cotton fiber development; the fiber length was determined by the elongation stage which was the most active period of biochemical reaction and critical to the fiber quality. The studies of the genetic law of cotton fiber showed that the genetic patterns of the same fiber-trait cotton differed in diverse genotypes. Quantitative trait loci (QTL) related to fiber quality and yield were distributed in the whole cotton genome. Some stable major QTL (such as FS1, qLI17 and qFL-Chr14-3) deserved further attention from researchers and were expected to be used in molecular aided selection application. Recent advances in the quality trait genes had clarified that Ligon lintless-1 (Li₁) mutant was an acting gene, and the N1 was a transcription factor named as MYB25-like. The deep sequencing in the fields of transcriptomics, proteomics and epigenetics effectively established the connection between RNA levels and protein levels, coding regions and non-coding regions, and found a series of genes and processes, such as transcription factors, lipid transfer protein encoding genes, genes involved in calcium signaling, genes related to polysaccharide synthesis, abundant miRNAs, and DNA methylation and so on, were all involved in cotton fiber development. This paper will provide a more comprehensively theoretical understanding for further research on the development mechanism and gene regulatory network of cotton fiber.
- plant genetics,
- cotton,
- fiber development,
- genomics,
- proteomics,
- epigenetics,
- functional genes,
- review

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棉纤维是由外珠被表皮层的单细胞分化发育而成，分为长绒(lint)和短绒(fuzz)2种，长绒是高级棉纱纺织品的主要原材料，短绒主要用做制作纤维素、絮棉、纸张及纺织品的原料。在已有的四倍体棉种中，陆地棉Gossypium hirsutum和海岛棉Gossypium barbadense已经被驯化为栽培种^[1-3]。目前世界上97%的棉纤维都产自陆地棉，产量高且适应性广，但是纤维品质中等；海岛棉产量低，适应性差，栽培范围不广泛，但是其纤维更长且品质高。如何获得优质高产的棉种，一直是遗传育种学家关注的焦点。而随着遗传学、细胞学和分子生物学等学科的交叉融合，棉纤维生长发育分子机制已成为国内外研究的热点。探明棉花种子表皮细胞生长发育的分子基础，对于提高棉花产量及改良纤维品质至关重要。早期有关棉纤维发育研究大多集中于遗传定位。大量与纤维品质和产量相关的数量性状位点(QTL)通过图位克隆的方法被发现于各个染色体^[4-5]，而光子显性基因Li₁，Li₂，N₁和Fbl以及光子隐性基因n₂，sma-4(fz)和sma-4(ha)^[6]等一直备受关注。近年来，深度测序技术的兴起，对棉纤维发育的分子机制的研究起了有效的推动作用。随着深度测序技术的不断革新，棉花全基因组测序不断完善^[7]，全基因组微卫星序列得以注释^[8]，单核苷酸多态性(SNP)芯片的开发成为可能^[9]，使得遗传定位工作更加便捷^[9-10]。转录组学、蛋白组学及表观遗传学领域三方位的深度测序有效构建了核糖核酸(RNA)水平和蛋白质水平、编码区域和非编码序列之间的联系，并发现一系列的转录因子、编码转脂蛋白的基因、钙信号转导相关基因、多糖合成相关蛋白、大量的微核糖核酸(miRNA)以及脱氧核糖核酸(DNA)甲基化作用等共同参与棉纤维发育过程^[11-16]。本文将从棉纤维发育各时期的形态结构变化及特征，经典遗传学研究，深度测序技术在转录组学、蛋白组学及表观遗传学领域的运用，以及棉纤维发育各个时期所涉及的相关调控基因等4个方面对棉纤维发育机制的研究进展进行综述。

1. 棉纤维发育各时期的形态结构变化特征

棉纤维细胞发育进程是一个多基因调控的有序的系统发生过程，整个细胞分化过程可被分为棉纤维起始、伸长、次生壁合成与增厚、脱水成熟等4个时期^[17-20]。

1.1. 棉纤维起始期

长绒纤维细胞一般在开花前或开花当天就开始突起，而短绒纤维的突起要稍迟几天，两者的分化过程基本相似^[21]。RAMSEY等^[22]通过观察开花前16 d到开花当天胚珠的亚显微结构，发现开花前16 d到前3 d表皮细胞无差异，说明纤维原始细胞的分化与突起晚于开花前3 d。而在开花前2~3 d纤维原始细胞受生长素(IAA)和赤霉素(GA₃)的刺激开始产生纤维^[11]，开花当天，纤维原始细胞的分化与突起已基本完成。棉花纤维原始细胞分化与突起多少决定种子表面纤维数量，从而决定了棉纤维的产量。

1.2. 棉纤维伸长期

研究发现一般只有25%~30%的棉花种子表皮细胞(约2万个)能正常突起伸长，形成成熟的纤维^[23-24]。棉纤维的伸长几乎和突起同时进行，从开花当天开始，发生在细胞壁膨胀过程，纤维的最终长度取决于纤维伸长速率和持续时间2个方面^[25]，一般持续20~30 d，该过程通过一种扩散生长机制实现并指导纤维细胞的极化生长^[26-27]。纤维伸长分为非极性膨胀和极性伸长2个阶段：非极性膨胀期决定了纤维的细度，纤维细胞向四周扩展直至形成纤维的最终直径^[28]；极性伸长期可使纤维长度达到最终长度的80%^[29]，这一时期生化反应最为活跃，主要决定纤维的长度，是影响纤维品质的关键时期^[24]。

1.3. 棉纤维次生壁合成与增厚期

棉纤维次生壁合成与增厚期和纤维伸长期存在一段时期的重叠^[17]，在开花后16 d开始，持续到开花后40 d，在这段时期，纤维素大量沉积，次生壁不断加厚^[30]。伴随着纤维素沉积的加速，纤维伸长逐渐减弱，该过程是影响纤维强度和韧性的关键时期。

1.4. 棉纤维脱水成熟期

在次生壁合成与增厚期后就进入了纤维脱水成熟期，发生在开花后40~50 d，棉铃开裂至充分吐絮，纤维失水，形成转曲^[31]。成熟的棉纤维由外向内依次为初生壁、次生壁和中腔。

2. 棉花纤维生长发育的遗传学研究

遗传规律研究和基因遗传定位是经典遗传学中2项重要的基础工作。在棉纤维遗传规律研究中发现，相同性状的材料基因型不同，其遗传模式也不同，而棉纤维发育相关基因的遗传定位又可被分成质量性状和数量性状(QTL)的定位。

2.1. 棉纤维遗传规律研究

CARVER^[32]和KEARNEY等^[33-34]研究发现棉花光子性状主要由2对独立的位点控制，显性光子基因(N₁)和隐性光子基因(n₂)，宋丽等^[35]证实这2种光子基因均符合单基因遗传模型。既无长绒也无短绒的L40突变体的光子性状为不完全显性^[36]。既无长绒也无短绒的突变体Xu142 fl的短绒的发育受N₁和n₂ 2对基因控制，长绒的发育受Li₃基因位点控制^[37]。陆地棉短绒突变体Li₁和Li₂均为单基因显性遗传^[38-40]。孙亚莉等^[41]选取大量的陆地棉和海岛棉的光子材料对棉花光子性状进行了遗传分析，其研究发现棉花短绒多少与生态环境有关系，且不同品种光子材料的遗传模式也不同，不论海岛棉还是陆地棉材料均存在显性、部分显性和隐性遗传。对3个陆地棉隐性性状的材料进一步研究表明：这3个材料的遗传规律均不同，‘库光子’的光子性状由2对隐性等位基因控制，并且有互补效应；‘陆无絮’的光子性状由2对隐性等位基因控制，基因间呈积加作用；SA65的光子性状由单隐性基因控制。

2.2. 棉纤维基因的QTL定位

纤维品质性状包括长度、整齐度、伸长率、强度、细度、颜色和马克隆值等多个方面。随着分子标记的不断开发与应用，在棉花染色体A组和D组染色体上都有大量棉纤维品质和产量相关的QTL被发现(表 1)。从表 1可知：纤维品质和产量性状的QTL几乎遍布了每一条染色体，且不同实验室使用不同的群体所得到的结果也有很大差异。同时，研究也发现这些性状受环境的影响很大，某些QTL在不同环境条件下有变化，甚至检测不到，导致已定位的QTL间重复性差^{[10, 42]}，这也说明纤维品质及产量性状的遗传非常复杂。研究也发现了一些稳定的主效QTL，如第10号染色体的棉纤维强度主效QTL(FS1)，解释了超过30.00%的表型变异^[43]；第19号染色体影响衣分的QTL(qLI17)，解释24.30%的表型变异^[34]；第8号染色体上颜色相关QTL(Ge6_Rd_8_3_10.60_[+])，解释48.00%的表型变异^[5]；以及第14号染色体上与长度相关的QTL(qFL-Chr14-3)，解释15.05%的表型变异^[10]，等等。此外，有些QTL虽然微效，但在不同环境下都能稳定存在，比如WANG等^[42]在8，11，12和21号染色上发现的6个QTL：qFL-A8-1(长度相关)，qFS-A8-1(强度相关)，qFS-A12-1(强度相关)，qFS-A12-2(强度相关)，qFS-D11-1(强度相关)和qFM-A11-1(马克隆值相关)。这些稳定存在的QTL都值得科研工作者进一步关注和研究。

性状	QTL所在染色体或连锁群	检出限(LOD)	变异率1%	群体	出处
长度	Chr04，Chrl8，Chr22	2.00~2.74	7.80~12.60	陆地棉TM-1×海岛棉3-79的F₂群体	KOHEL等^[44]
	Chr20，LGA02(Chr08)，LGA03 (Chr11)，LGA05	2.63~5.40	2.90~13.70	陆地棉Siv’ on×海岛棉F-177的F₂和F₃群体	PATERSON等^[4]
	Chr04	3.50	24.00	陆地棉Acala 44×海岛棉Pima S-7的F₂群体	MEI等^[45]
	Chr01，Chr03，Chr04，Chr06，Chr09，Chr13，Chr14，Chr18，Chr19，Chr20，Chr21，Chr23，Chr24，Chr26	3.30~9.50	6.00~40.00	陆地棉Guazuncho-2×海岛棉VH8-4602RIL	LACAPE等^[5]
	Chr05，Chr07，Chr08，Chr11，Chr12，Chr19，Chr21，Chr23，Chr26	4.57~6.05	2.47~8.49	陆地棉TM-1×海岛棉Hai7124的CSILs群体	WANG等^[42]
	Chr10，Chr14，Chr15	2.50~7.71	6.21~15.05	陆地棉HS46 ×陆地棉MAR CABU-CAG8US-1-88 RIL	LI等^[10]
整齐度	Chr04，Chr14，Chr15，Chr22，LGA03 (Chr11)，LGA05	1.65~3.79	2.10~13.30	陆地棉Siv’on×海岛棉F-177的F₂和F₃群0体	PATERSON等^[4]
	Chr05，Chr09，Chr12，Chr15，Chr16，Chr18，Chr19，Chr20，Chr23，Chr26	3.50~7.80	9.00~32.00	陆地棉Guazuncho-2×海岛棉VH8-4602 RIL	LACAPE等^[5]
	Chr09	2.68~4.17	5.58~10.94	陆地棉HS46 ×陆地棉MARCABU- CAG8US-1-88的RIL	LI等^[10]
伸长率	Chr05，Chr10，Chr15，Chr23，LGA02 (Chr8)，LGA03(Chr11)，LGD07	2.32~5.77	3.40~8.90	陆地棉Siv’on×海岛棉F-l77的F₂和F₃群体	PATERSON等^[4]
	Chr09	5.16	42.00	陆地棉Acala 44×海岛棉Pima S-7的F₂群体	MEI等^[45]
	Chr02，Chr06，Chr09，Chr10，Chr12，Chr13，Chr15，Chr19，Chr20，Chr21，Chr23，Chr26	3.40~6.70	6.00~21.00	陆地棉Guazuncho-2×海岛棉VH8-4602RIL	LACAPE等^[5]
	Chr14，Chr20，Chr24	2.49~7.80	5.35~32.28	陆地棉HS46 ×陆地棉MARCABU-CAG8US-1-88 RIL	LI等^[10]
强度	Chr03，Chr14，Chr15，Chr25	2.08~2.69	10.40~23.10	陆地棉TM-1×海岛棉3-79的F₂群体	KOHEL等^[10]
	Chr10	4.79~5.80	53.00~53.80	异质棉7235×陆地棉TM-1的F₂群体	ZHANG等^[43]
	Chr01，Chr04，Chr14，Chr17，Chr18，Chr20，Chr22，Chr23，Chr25，LGA01 (Chr13)，LGA02(Chr08)，LGA03(Chr11)，LGA05，LGD02(Chr21)，LGD03(Chr24)，LGD04，LGD07	0.21~6.22	2.50~17.40	陆地棉Siv’on×海岛棉F-177的F₂和F₃群体	PATERSON等^[4]
	Chr03，Chr04，Chr05，Chr07，Chr09，Chr12，Chr14，Chr15，Chr16，Chr18，Chr19，Chr21，Chr23，Chr26	3.30~8.50	7.00~31.00	陆地棉Guazuncho-2×海岛棉VH8-4602RIL	LACAPE等^[5]
	Chr05，Chr07，Chr08，Chr09，Chr11，Chr12，Chr13，Chr14，Chr15，Chr16，Chr17，Chr18，Chr21，Chr23	7.32~22.54	5.07~15.82	陆地棉TM-1×海岛棉Hai7124的CSILs群体	WANG等^[42]
细度	Chr01，Chr02，Chr03，Chr12，Chr16，LGD01	2.16~4.04	16.70~43.90	陆地棉TM-1×海岛棉3-79的F₂群体	KOHEL等^[44]
	Chr02，Chr04，Chr05，Chr06，Chr09，Chr14，Chr15，Chr17，Chr20，Chr23，Chr25，LGA01 (Chr13)，LGA05，LGA06，LGD01，LGD02(Chr2l)，LGD03(Chr24)，LGD04，LGD05，LGD07	2.21~9.78	2.20~30.30	陆地棉Siv’on×海岛棉F-l77的F₂和F₃群体	PATERSON等^[4]
	Not determined	5.11	43.20	陆地棉Acala 44×海岛棉Pima S-7的F₂群体	MEI等^[45]
	Chr01，Chr02，Chr03，Chr04，Chr05，Chr06，Chr08，Chr09，Chr10，Chr12，Chr15，Chr16，Chr17，Chr18，Chr19，Chr20，Chr21，Chr22，Chr23，Chr24，Chr25，Chr26	3.30~8.90	6.00~41.00	陆地棉Guazuncho-2×海岛棉VH8-4602RIL	LACAPE等^[5]
颜色	Chr06，Chr09，Chr14，Chr17，Chr18，Chr22，Chr25，LGA01，LGA02，LGA03，LGD02(Chr21)	2.66~11.67	2.50~14.90	陆地棉Siv’on×海岛棉F-177的F₂和F₃群体	PATERSON等^[4]
	Chr01，Chr02，Chr06，Chr07，Chr08，Chr09，Chr11，Chr14，Chr15，Chr17，Chr18，Chr19，Chr21，Chr22，Chr25	3.30~10.60	6.00~48.00	陆地棉Guazuncho-2×海岛棉VH8-4602RIL	LACAPE等^[5]
马克隆值	Chr05，Chr06，Chr09，Chr11，Chr12，Chr15，Chr16，Chr19，Chr21，Chr22	4.56~9.09	0.80~8.03	陆地棉TM-1×海岛棉Hai7124的CSILs群体	WANG等^[42]
	Chr14，Chr16	2.51~4.23	5.52~9.20	陆地棉HS46 ×陆地棉MARCABU- CAG8US-1-88 RIL	LI等^[10]
产量	A02(Chr08)，A03(Chr11)，Chr14，Chr23，Chr25，LG5	3.00~5.28	13.01~28.35	陆地棉Handan 208×海岛棉Pima 90的F₂群体	HE等^[46]
衣分	D08(Chr19)	3.45	24.34	陆地棉Handan 208×海岛棉Pima 90的F₂群体	HE等^[46]

Table 1. QTL related to cotton fiber quality and yield in different populations

然而，棉纤维相关性状QTL的分离和克隆仍然很少。有研究发现在第12条染色体上(A12/D12)与棉纤维品质相关的QTL附近的GhHOX3基因对棉纤维长度起重要调控作用^[47]，以及定位于同源染色体A8(chr08)和D8(chr24)上的GhSusA1基因过表达可以增强纤维长度和强度^[48]。

2.3. 棉纤维基因的质量性状定位

在光子显性基因定位中发现，N₁和Fbl基因位于chr12上^[6]，其中N₁基因被鉴定为转录因子MYB25-like^[49]。光子隐性基因定位研究表明：n₂定位在chr26上，sma-4(fz)位于L.G.A3的端部，sma-4(ha)位于L.G.A3中部^[6]。超短纤维突变体Li₁基因位于chr22上，并已通过精细定位被克隆到，是一个肌动蛋白家族基因^[50-53]；而Li₂基因则位于chr18上^{[6, 53]}。

3. 棉纤维发育组学研究

棉纤维发育过程涉及到大量的基因和通路调控。利用高通量测序技术，对棉纤维发育的转录组和蛋白组分析及表观遗传研究，可以短时间内获得大量信息，捕获到许多参与不同发育阶段的特异性基因及信号通路，为下游单独研究重要基因的功能奠定良好的基础。

3.1. 转录组学研究

利用棉花胚珠体外培养技术结合转录组数据分析比较，KIM等^[11]在纤维起始分化时期发现了许多在野生型和无毛突变体间表达有差异的基因，包括MYB25，MYB109，PDF1，MYB25-like，HD1等转录调节因子，表明在棉纤维起始分化过程中存在复杂的信号网络调节机制。通过比较短绒突变体(Li₁)与野生型之间在开花后1，3和8 d的转录组数据，LIANG等^[49]在胚珠中共检测到7 852个差异表达基因，主要参与次生代谢物和脂质代谢途径，其中涉及非长链脂肪酸生物合成的37个基因在Li₁突变体纤维的快速伸长发育过程中被显著抑制，这说明脂质代谢途径与纤维伸长密切相关。HOVAV等^[54]评估了从开花后初级次生壁到次级次生壁合成过程中棉纤维发育的转录组变化发现，棉纤维发育过程中的基因转录水平很高，在每个阶段占到所有基因的75%~94%，并且半数以上的基因在纤维发育的至少一个阶段中上调。BOLTON等^[55]利用基因芯片技术和实时荧光定量PCR技术在Li₁突变体中发现超过100个基因在次生壁的生物合成过程中差异表达，其中的3个候选基因：伸展蛋白(extensin)，蔗糖合成酶(sucrose synthase)和微管蛋白(actin)的表达量明显偏离野生型的表达水平。通过陆地棉TM-1背景下的海岛棉染色体导入系与亲本纤维的转录组差异比较，FANG等^[56]在CSIL-35431和CSIL-31010等2个导入系的次生壁合成过程中发现了大量与TM-1有表达差异的基因，功能富集分析表明这些基因主要富集于次生细胞壁的生物合成、葡糖醛酸合成、纤维素合成等生物途径。

3.2. 蛋白质组学研究

利用蛋白组学研究，许多棉纤维发育过程中的重要蛋白被不断发掘，且此技术很好地互补了转录组只能在mRNA水平上研究纤维相关基因的劣势。HU等^[12]应用相对和绝对定量(iTRAQ)LC-MS/MS分析技术研究了1 317个纤维特异性表达蛋白，其中205个蛋白在发育阶段中差异表达，190个蛋白在野生和栽培棉之间差异表达。结合转录组、iTRAQ蛋白质组和遗传图谱定位的综合分析方法，MA等^[13]发现徐州142野生型与其无绒毛突变体(fl)的胚珠之间存在大量差异表达的基因和蛋白，这些差异基因和蛋白主要存在于氨基酸、核苷酸、脂肪酸和叶酸代谢以及黄酮生物合成中，说明这些代谢途径在纤维发育过程中具有重要作用。

3.3. 表观遗传学研究

近年来，棉纤维发育的表观遗传学研究也取得了巨大进展。基于pre-miRNAs和已发现的miRNA靶基因数据，CHEN等^[14]对83个miRNA前体及其目标调控基因进行了研究，并构建了miRNAs及其靶位点调控网络，并揭示了这些miRNA及靶基因在纤维不同发育阶段的表达模式。SONG等^[15]对纤维和胚珠进行了甲基化组、转录组和小RNA组学分析，发现在胚珠和纤维发育过程中CHH甲基化变化显著。该研究发现，在胚珠中，启动子中的CHH甲基化与可诱导RNA依赖的DNA甲基化(RdDM)和胚珠偏好基因上调的siRNA呈正相关；在纤维细胞中，胚珠衍生细胞产生独立的RdDM的异染色质CHH超甲基化，抑制转座子及附近纤维相关基因的活性。使用甲基化抑制剂5-氮-2′-脱氧胞苷对胚珠进行体外培养，可以减少纤维细胞的数量和长度，这表明DNA甲基化在纤维发育中具有潜在作用。这些研究表明：启动子和转座子及附近基因中的RdDM依赖的甲基化可作为基因和转座子表达的双保险反馈机制。ZOU等^[16]对参与纤维起始和伸长过程的长链非编码RNA(lncRNA)进行了系统分析，共鉴定到5 996条lncRNAs，其中长链非编码RNA(lincRNA)3 510条，天然反义转录RNA(lncNAT)2 486条，表明lncRNA对棉纤维的发育至关重要。

4. 棉纤维发育不同时期相关调控基因研究

4.1. 棉纤维起始期相关基因

在棉花胚珠EST数据库中，约10%的基因与转录因子密切相关，包括56个转录因子家族成员^[57]，如MYB类转录因子家族的GL1及其同源基因MYB2^[58]，MYB109^[59]，TTG1^[60]和GL2^[61]等均被发现在棉纤维发育的早期阶段高效表达。棉花GaMYB2基因可以诱导种子表皮毛的产生，且转化拟南芥可以弥补GL1突变对表皮毛起始分化造成的影响^[58]。通过干扰GhMYB109的表达，PU等^[62]发现棉纤维细胞分化延迟且起始数量减少，说明GhMYB109在棉花纤维分化阶段起重要作用。LOGUERICO等^[63]发现GhMYB4和GhMYB5基因在纤维分化期的胚珠中特异表达。WALFORD等^[64]发现GhHD1可以介导棉花表皮细胞的分化。辛婧^[65]的研究表明转录因子GbSPB8可能调控棉花纤维起始发育。

4.2. 棉纤维伸长期相关基因

转脂蛋白和钙信号转导相关蛋白在棉纤维伸长过程中起到极其重要的作用。MA等^[66]研究发现：转脂蛋白GhLTP3和GhLTP6的表达量在纤维快速伸长期达到最高水平，此外李锡花等^[67]发现GhLTP3的表达量从开花后0~15 d中表达量不断升高，在第15天达到顶峰，之后逐渐下降。赵存鹏等^[68]和GAPPER等^[69]研究发现：钙调蛋白CaM在低温逆境的条件下能使活性氧(ROS)、超氧游离基、过氧化氢和羟自由基等物质提高，进而使纤维细胞壁松弛，从而影响细胞伸长。CHENG等^[70]发现GhCaM7-like基因在纤维快速伸长期显著表达。HUANG等^[71]发现钙依赖性蛋白激酶GhCPK1在开花第10天的胚珠中高表达。这些研究都说明钙信号转导在纤维伸长过程中发挥重要作用。除了转脂蛋白和钙信号转导相关蛋白，转录调节因子也参与其中，如ZHANG等^[72]发现GbMYB25与GbML1相互作用并通过调节ROS信号调节纤维伸长。HSU等^[73]发现GhMYB7可以调控LTP3等脂转移蛋白编码基因。

4.3. 棉纤维次生壁合成与加厚期相关基因

DELMER等^[74]发现Rac9和Rac13可以控制棉花纤维素沉积方向。ZHAO等^[75]发现GhRGP1在纤维发育的初生壁伸长及次生壁加厚后期优势表达，参与植物细胞壁非纤维素类的多糖合成。杨郁文等^[76]发现一种Ser/Thr激酶和Try激酶的双受体蛋白GhRLK1与激活和维持次级细胞壁形成的细胞信号传导过程有关。此外，GhRDL1(RD22-Like1)与GhEXPA1互作会影响纤维细胞壁发育，GhRDL1基因过表达会产生长且质量较好的棉纤维^[77]。

4.4. 棉纤维脱水成熟期相关基因

目前，关于棉纤维细胞脱水成熟期的相关研究较少，对于纤维在脱水成熟过程中细胞与分子水平的变化还不清楚，只是猜想可能涉及到棉纤维细胞的程序性死亡^[78]。

5. 研究展望

随着技术的革新，大量棉纤维发育相关基因和涉及的调控网络被不断发现，但是基因间的相互作用及其潜在的调控机制还有待进一步探索。近年来，棉花转基因技术日益成熟，新兴的CRISPR/Cas9基因编辑系统也于2017年3月在棉花基因组靶基因敲除中得到首次应用。JANGA等^[79]利用CRISPR/Cas9系统成功将转绿色荧光蛋白(GFP)基因棉花系的GFP基因敲除；LI等^[80]以棉花內源GhMYB25为目标基因，使用2种单导向RNA对该基因进行定点突变，突变率分别为100%和98.8%。这些研究表明：CRISPR/Cas9可以在棉花基因组上进行高效和高特异性地突变。随着新技术在棉花中的应用与成熟，棉纤维发育相关基因及其调控机制也有望有更多的发现。

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Genetic characteristics and research advances of genes related to cotton fiber development

doi: 10.11833/j.issn.2095-0756.2018.06.021