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作物学报 ›› 2026, Vol. 52 ›› Issue (6): 1682-1697.doi: 10.3724/SP.J.1006.2026.54138

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

陆地棉I类LBD家族成员鉴定及GhLBD6调控开花期的功能和单倍型分析

彭佳泺1(), 李颖1, 李丹丹1, 杨军宁1, 郭学峰1, 张文姣1, 俞晓雪1, 周亚荣1, 王振玉2, 王彩香1, 马雄风2,*(), 宿俊吉1,*()   

  1. 1 甘肃农业大学生命科学技术学院, 甘肃兰州 730070
    2 中国农业科学院棉花研究所, 河南安阳 455000
  • 收稿日期:2025-11-05 接受日期:2026-02-27 出版日期:2026-06-12 网络出版日期:2026-03-09
  • 通讯作者: * 宿俊吉, E-mail: sujj@gsau.edu.cn; 马雄风, E-mail: maxiongfeng@caas.cn
  • 作者简介:彭佳泺, E-mail: 3028627812@qq.com
  • 基金资助:
    国家自然科学基金项目(32572431);新疆自治区重点研发任务专项项目(2025B02001-2);财政部和农业农村部国家现代农业产业技术体系建设专项(Cotton, CARS-15-07);棉花生物育种与综合利用全国重点实验室开放课题基金项目(CB2024A08)

Identification of class I LBD family members in upland cotton and function and haplotype analyses of GhLBD6 in regulating flowering period

Peng Jia-Luo1(), Li Ying1, Li Dan-Dan1, Yang Jun-Ning1, Guo Xue-Feng1, Zhang Wen-Jiao1, Yu Xiao-Xue1, Zhou Ya-Rong1, Wang Zhen-Yu2, Wang Cai-Xiang1, Ma Xiong-Feng2,*(), Su Jun-Ji1,*()   

  1. 1 College of Life Science and Technology, Gansu Agricultural University, Lanzhou 730070, Gansu, China
    2 Institute of Cotton Research of Chinese Academy of Agricultural Sciences, Anyang 455000, Henan, China
  • Received:2025-11-05 Accepted:2026-02-27 Published:2026-06-12 Published online:2026-03-09
  • Contact: * Su Jun-Ji, E-mail: sujj@gsau.edu.cn; Ma Xiong-Feng, E-mail: maxiongfeng@caas.cn
  • Supported by:
    National Natural Science Foundation of China(32572431);Key Research and Development Task Project of Xinjiang Uygur Autonomous Region(2025B02001-2);China Agriculture Research System of MOF and MARA(Cotton, CARS-15-07);State Key Laboratory of Cotton Bio-breeding and Integrated Utilization Open Fund(CB2024A08)

摘要:

侧生器官边界域(lateral organ boundaries domain, LBD)家族基因在植物多种生物学过程中发挥重要作用。前人研究表明, I类LBD基因参与调控植物开花时间, 然而陆地棉I类LBD基因是否具有类似功能尚不清楚。本研究在全基因组水平鉴定并分析陆地棉I类LBD家族成员; 通过单倍型分析筛选调控开花相关候选基因; 利用RNA-seq及RT-qPCR确定候选基因在不同组织和早晚花品种中的表达模式; 通过病毒诱导的基因沉默(virus-induced gene silencing, VIGS)验证目标基因的功能, 并对其进行育种演化分析及分子标记开发。结果显示, 在陆地棉中鉴定出102个I类GhLBDs基因, 这些基因不均匀分布在26条染色体上, 按照进化关系分为6个组别, 同一组别内的LBD成员具有相似的基序组成与排列模式。通过GhLBDs的单倍型与表达分析, 筛选出开花期相关候选基因GhLBD6。通过VIGS技术结合石蜡切片、荧光定量等, 发现抑制GhLBD6表达会导致植株花芽分化提前, 现蕾期和开花期分别显著提前6.57 d和6.86 d。另外, GhLBD6编码区存在2种单倍型: GhLBD6-Hap 1和GhLBD6-Hap 2。其中, GhLBD6-Hap 1为早花的优异等位变异, 在陆地棉育种过程中受到人工选择。最后开发了可用于区分GhLBD6两种单倍型的竞争性等位基因特异性PCR (kompetitive allele-specific PCR, KASP)分子标记。本研究结合正反向遗传学试验结果, 证明了GhLBD6参与陆地棉开花期调控, 为棉花早熟分子育种奠定了基础。

关键词: 陆地棉, GhLBD6, 开花期, 单倍型, 竞争性等位基因特异性PCR分子标记

Abstract:

Lateral organ boundaries domain (LBD) family genes play important roles in a wide range of plant biological processes. Previous studies have shown that class I LBD genes are involved in the regulation of flowering time; however, whether class I LBD genes have similar functions in upland cotton remains unclear. In this study, we identified and characterized class I LBD family members at the whole-genome level in upland cotton and screened flowering-related candidate genes through haplotype analysis. The expression patterns of candidate genes in different tissues and in early- and late-flowering varieties were examined using RNA-seq and RT-qPCR, respectively. The function of the target gene was validated by virus-induced gene silencing (VIGS), followed by analysis of its breeding-related evolutionary dynamics and the development of molecular markers. In total, 102 class I GhLBD genes were identified; they were unevenly distributed across 26 chromosomes and were classified into six groups based on evolutionary relationships. LBD members within the same group showed similar motif composition and arrangement. Integrating haplotype and expression analyses, GhLBD6 was identified as a candidate gene associated with flowering time. VIGS combined with paraffin sectioning and RT-qPCR showed that suppression of GhLBD6 accelerated floral bud differentiation and significantly advanced budding and flowering by 6.57 d and 6.86 d, respectively. Two haplotypes (GhLBD6-Hap 1 and GhLBD6-Hap 2) were identified in the GhLBD6 coding region; among them, GhLBD6-Hap 1 represents a favorable early-flowering allele and appears to have been under artificial selection during upland cotton breeding. Finally, we developed a kompetitive allele-specific PCR (KASP) molecular marker to distinguish the two GhLBD6 haplotypes. Together, these forward- and reverse-genetic results demonstrate that GhLBD6 contributes to flowering-time regulation in upland cotton, providing a basis for molecular breeding of early-maturing cotton.

Key words: upland cotton, GhLBD6, flowering period, haplotype, KASP molecular marker

附表1

本研究所用引物列表"

引物名称
Primer name
序列
Sequence (5′-3′)
引物用途
Primer usage
GhLBD6-F GTTCACAAGGTGTTCGGTGC 克隆沉默片段和验证单倍型
Clone silent segments and verify haplotypes
GhLBD6-R GACGTTCATGGCGAGGAGAT
GhLBD6-MF G GAATTCGTTCACAAGGTGTTCGGTGC 克隆带酶切位点的沉默片段
Clone silent segments with enzyme-cutting sites
GhLBD6-MR GG GGTACCGACGTTCATGGCGAGGAGAT
GhLBD6-QF GATCTCAGCTGTGCTAAGTCG RT-qPCR (GH_A11G3479)
GhLBD6-QR TCCTCCTCCTCCACCACT
GhLBD20-2-QF CAACAGCAACACTTAGCATCG RT-qPCR (GH_A13G0147)
GhLBD20-2-QR CTTCGTCGTCATGACACGG
GhLBD12-1-QF AATGCAGCACAACCAGGAG RT-qPCR (GH_A10G2646)
GhLBD12-1-QR TGCAGCAGGGTCTTGAATTAC
GhActin-QF ATCCTCCGTCTTGACCTTG RT-qPCR (内参基因)
RT-qPCR (endogenous gene)
GhActin-QR TGTCCGTCAGGCAACTCAT
GhAP1-QF ATGTGGAGCAGGCACATTGG RT-qPCR (GH_D03G1119)
GhAP1-QR TCATCCATGGCAGCAAGACG
GhSOC1-QF AAGTACGGTCTGTTGCCAGG RT-qPCR (GH_A11G0861)
GhSOC1-QR CTGTTGAGTTGGGTTGCACG
GhSVP-QF ATTGGGTTGAGCCGTGTGAT RT-qPCR (GH_D12G0863)
GhSVP-QR GGTAACTTGTCTTGGGCCGT
GhLFY-QF TGGCAAAAAGAACGGCCTTG RT-qPCR (GH_D07G0531)
GhLFY-QR TAGTGGGGCATTTTTCGCCA
GhCAL-QF CGACGGAACATGCACAACAT RT-qPCR (GH_A07G0875)
GhCAL-QR GGCTTCGTAACTCCCTCCAA
1-FAM GAAGGTGACCAAGTTCATGCTTCCTCCTCCTCCACCACC 基因分型Genotyping
2-HEX GAAGGTCGGAGTCAACGGATTTCCTCCTCCTCCACCACT
Reverse primer GATCTCAGCTGTGCTAAGTCG

附表2

9物种I类LBD家族成员的鉴定"

物种
species
1组
Group 1
2组
Group 2
3组
Group 3
4组
Group 4
5组
Group 5
6组
Group 6
总计
Total
拟南芥Arabidopsis thaliana 2 3 3 8 10 5 31
陆地棉Gossypium hirsutum 12 18 12 15 32 13 102
亚洲棉Gossypium arboreum 6 8 6 8 17 5 50
雷蒙德氏棉Gossypium raimondii 6 9 6 7 17 6 51
苹果Malus domestica 4 7 6 9 11 10 47
水稻Oryza sativa 1 2 3 3 8 6 23
玉米Zea mays 4 5 4 5 11 7 36
大麦Hordeum vulgare 2 3 3 4 7 4 23
小麦Triticum aestivum 6 9 9 9 15 12 60
总计Total 43 64 52 68 128 68 423

图1

I类LBD家族成员的鉴定和系统发育分析 A: 9个物种I类LBD家族成员的系统发育进化树; B: I类LBD家族成员在不同组别的数目分布。"

附表3

陆地棉102个I类LBD基因的基本信息"

基因ID
Gene ID
重命名
Rename
分组
Group
开花期相关SNP显著情况
Significance of SNP related to flowering period
2种主要单倍型样本数比值
Sample size ratio of two major
haplotypes
GH_D07G1252 GhLBD1-4 1
GH_A07G1269 GhLBD11-3 1
GH_A08G0118 GhLBD11-1 1
GH_D08G0123 GhLBD11-2 1 P = 8.389E-03 33/246
GH_A13G0031 GhLBD1-1 1
GH_D13G0027 GhLBD11-4 1
GH_A09G1214 GhLBD1-7 1
GH_D09G1171 GhLBD1-8 1
GH_A06G1943 GhLBD1-2 1
GH_D06G1967 GhLBD1-3 1
GH_D04G0866 GhLBD1-5 1
GH_A05G3610 GhLBD1-6 1
GH_A05G1095 GhLBD12-9 2
GH_D05G1086 GhLBD12-10 2
GH_D10G2744 GhLBD12-5 2 P = 1.752E-03 85/330
GH_A10G2639 GhLBD12-6 2
GH_D06G1896 GhLBD12-7 2
GH_A06G1863 GhLBD12-8 2
GH_D03G1614 GhLBD12-1 2 P = 2.065E-04 92/138
GH_A03G0359 GhLBD12-2 2
GH_D08G2310 GhLBD12-3 2
GH_A08G2303 GhLBD12-4 2
GH_A01G0920 GhLBD4-7 2
GH_D01G0938 GhLBD4-8 2
GH_D10G0505 GhLBD4-2 2 P = 5.429E-03 49/368
GH_A10G0479 GhLBD4-3 2
GH_A06G1622 GhLBD4-1 2 P = 2.113E-05 35/365
GH_D06G1660 GhLBD4-4 2 P = 0.013 66/335
GH_A05G1451 GhLBD4-5 2
GH_D05G1469 GhLBD4-6 2
GH_D05G2831 GhLBD15-1 3 P = 3.016E-04 61/339
GH_A05G2818 GhLBD15-2 3
GH_A05G4224 GhLBD15-3 3
GH_D04G0152 GhLBD15-4 3
GH_A13G0058 GhLBD15-7 3
GH_D13G0055 GhLBD15-8 3
GH_D03G1635 GhLBD15-5 3
GH_A03G0337 GhLBD15-6 3
GH_D13G0154 GhLBD21-1 3
GH_A13G0147 GhLBD21-2 3
GH_A10G2646 GhLBD21-3 3
GH_D10G2751 GhLBD21-4 3
GH_A11G1984 GhLBD36-1 4
GH_D11G2021 GhLBD36-2 4
GH_D12G0713 GhLBD36-5 4
GH_A12G0708 GhLBD36-6 4
GH_A10G0932 GhLBD36-3 4
GH_D10G1030 GhLBD36-4 4
GH_A11G3479 GhLBD6 4 P = 3.446E-03 150/256
GH_A05G3760 GhLBD25-1 4
GH_D04G0587 GhLBD25-2 4
GH_A09G0336 GhLBD25-3 4
GH_D09G0343 GhLBD25-4 4
GH_D05G2568 GhLOB-1 4
GH_A05G2547 GhLOB-2 4
GH_A11G0685 GhLOB-3 4
GH_D11G0717 GhLOB-4 4
GH_D11G1053 GhLBD11-5 5
GH_A08G1506 GhLBD1-9 5
GH_D08G1519 GhLBD1-10 5
GH_D08G1617 GhLBD18-1 5
GH_A08G1603 GhLBD18-2 5
GH_D12G1185 GhLBD18-3 5
GH_A12G1184 GhLBD18-4 5
GH_D11G0877 GhLBD19-1 5
GH_A11G0839 GhLBD19-2 5
GH_A12G1183 GhLBD31-1 5
GH_D12G1183 GhLBD31-2 5
GH_D11G3430 GhLBD20-1 5
GH_A11G3425 GhLBD20-2 5 P = 0.0141 173/197
GH_A10G2454 GhLBD33-1 5 P = 6.662E-03 71/332
GH_D10G2563 GhLBD33-2 5
GH_A07G2389 GhLBD16-1 5
GH_D07G2331 GhLBD16-4 5 P = 1.482E-03 27/383
GH_D11G1372 GhLBD16-2 5
GH_A11G1341 GhLBD16-3 5
GH_D08G0672 GhLBD16-5 5
GH_D08G2179 GhLBD16-6 5
GH_A08G2159 GhLBD16-7 5
GH_D08G2180 GhLBD29-6 5
GH_A08G2160 GhLBD29-7 5
GH_A11G1342 GhLBD29-1 5 P = 0.0467 67/345
GH_D11G1373 GhLBD29-2 5
GH_A07G2390 GhLBD29-9 5
GH_D07G2332 GhLBD29-10 5 P = 3.417E-04 99/307
GH_D08G0670 GhLBD29-8 5
GH_A08G0669 GhLBD29-3 5
GH_D08G0671 GhLBD29-4 5
GH_A08G0670 GhLBD29-5 5 P = 0.0299 130/278
GH_A05G1199 GhLBD24-1 6 P = 1.708E-03 74/334
GH_D05G1201 GhLBD24-4 6 P = 2.748E-03 68/336
GH_A12G2803 GhLBD24-2 6
GH_D12G2831 GhLBD24-3 6 P = 3.667E-04 93/294
GH_A11G0311 GhLBD27-1 6
GH_D11G0325 GhLBD27-2 6
GH_A07G0225 GhLBD22-4 6
GH_D07G0232 GhLBD22-6 6
GH_D10G0628 GhLBD22-1 6
GH_A10G0592 GhLBD22-2 6
GH_D06G2227 GhLBD22-5 6
GH_A06G2192 GhLBD22-3 6
GH_D06G2226 GhLBD22-7 6

图2

陆地棉I类LBD家族成员的系统发育、基因和蛋白结构分析 A: 系统进化树; B: 保守基序分布; C: 保守结构域; D: 基因结构。CDS: 编码序列。"

图3

陆地棉I类LBD基因的染色体分布与共线性分析 A: I类LBD基因的染色体分布, A01-A13分别代表陆地棉At亚基因组上的13条染色体, D01-D13分别代表陆地棉Dt亚基因组上的13条染色体; B: 陆地棉(Gh)、亚洲棉(Ga)和雷蒙德氏棉(Gr)之间的种间共线性; C: 陆地棉种内共线性; D: 共线性基因对的Ka、Ks及Ka/Ks值分布。"

图4

陆地棉I类LBD家族成员的单倍型分析及候选基因表达分析 A: 自然变异位点分析; B: 单倍型分析, 按照2种主要单倍型样本数比值(较小样本数/较大样本数) ≥ 0.5筛选候选基因。黑色星星代表未选中的基因, 红色星星代表候选基因; C: 组织表达热图, 数值代表log2 (FPKM+1)。DPA代表开花后天数; D: 候选基因在早晚花品种中的相对表达。**表示在0.01水平差异显著; ns表示差异不显著。"

图5

陆地棉GhLBD6的VIGS沉默植株表型观察统计 A: 阳性对照的白化表型; B: GhLBD6的相对表达量, TRV:00和TRV:GhLBD6分别代表阴性对照和GhLBD6沉默植株; C: 三叶期顶端分生组织石蜡切片; D: 现蕾、开花表型; E: 开花相关基因的相对表达量。*和**分别表示在0.05和0.01水平差异显著; ns表示差异不显著。"

图6

GhLBD6的自然变异、单倍型及其与开花期的关联分析 A: GhLBD6编码区的单倍型组成; B: 单倍型与开花期关联分析; C: 单倍型验证; D: GhLBD6表达水平与开花期的线性回归分析; E: 不同单倍型群体中GhLBD6的表达水平。*和**分别表示在0.05和0.01水平差异显著。"

图7

GhLBD6优异等位变异育种演化分析及基因分型 A: 单倍型在不同地域陆地棉群体中的分布; B: 单倍型在不同年代陆地棉群体中的分布; C: 单倍型在极端早晚花品种中的频率分布; D: 核苷酸多样性的比值(Min-50/Max-50); E: 遗传分化指数; F: 等位基因鉴别图。YZRR: 长江流域棉区; YRR: 黄河流域棉区; NIR: 西北内陆棉区。不同小写字母表示在P < 0.05水平差异显著。"

[1] Shuai B, Reynaga-Peña C G, Springer P S. The LATERAL ORGAN BOUNDARIES gene defines a novel, plant-specific gene family. Plant Physiol, 2002, 129: 747-761.
[2] Iwakawa H, Ueno Y, Semiarti E, et al. The ASYMMETRIC LEAVES2 gene of Arabidopsis thaliana, required for formation of a symmetric flat leaf lamina, encodes a member of a novel family of proteins characterized by cysteine repeats and a leucine zipper. Plant Cell Physiol, 2002, 43: 467-478.
doi: 10.1093/pcp/pcf077
[3] Semiarti E, Ueno Y, Tsukaya H, et al. The ASYMMETRIC LEAVES2 gene of Arabidopsis thaliana regulates formation of a symmetric lamina, establishment of venation and repression of meristem-related homeobox genes in leaves. Development, 2001, 128: 1771-1783.
doi: 10.1242/dev.128.10.1771 pmid: 11311158
[4] Rong M, Gao S X, Wen D, et al. The LOB domain protein, a novel transcription factor with multiple functions: a review. Plant Physiol Biochem, 2024, 214: 108922.
doi: 10.1016/j.plaphy.2024.108922
[5] Okushima Y, Fukaki H, Onoda M, et al. ARF7 and ARF19 regulate lateral root formation via direct activation of LBD/ASL genes in Arabidopsis. Plant Cell, 2007, 19: 118-130.
doi: 10.1105/tpc.106.047761 pmid: 17259263
[6] Nguyen T H, Kim M J, Kim J. Transcription factors WRKY2 and WRKY34 control LATERAL ORGAN BOUNDARIES DOMAIN10 expression in pollen vegetative cell nuclei. Plant Physiol, 2024, 196: 2463-2475.
doi: 10.1093/plphys/kiae448
[7] Teng R D, Wu Z, Xu S J, et al. A novel lateral organ boundary-domain factor CmLBD2 positively regulates pollen development by activating CmACOS5 in Chrysanthemum morifolium. Plant Cell Physiol, 2021, 62: 1687-1701.
doi: 10.1093/pcp/pcab124
[8] Dang T V T, Lee S, Cho H, et al. The LBD11-ROS feedback regulatory loop modulates vascular cambium proliferation and secondary growth in Arabidopsis. Mol Plant, 2023, 16: 1131-1145.
doi: 10.1016/j.molp.2023.05.010
[9] Dong J H, Wang Y, Xu L, et al. RsLBD3 regulates the secondary growth of taproot by integrating auxin and cytokinin signaling in radish (Raphanus sativus L.). J Integr Plant Biol, 2025, 67: 1823-1842.
doi: 10.1111/jipb.v67.7
[10] Castaings L, Marchive C, Meyer C, et al. Nitrogen signalling in Arabidopsis: how to obtain insights into a complex signalling network. J Exp Bot, 2011, 62: 1391-1397.
doi: 10.1093/jxb/erq375 pmid: 21118821
[11] Zhang J L, Dong D H, Jia C Y, et al. Fine-tuning of MYC2- mediated Botrytis defense response by the LBD40/42-CRL3BPM4 module in tomato. Plant Cell, 2025, 37: koaf258.
doi: 10.1093/plcell/koaf258
[12] Srikanth A, Schmid M. Regulation of flowering time: all roads lead to Rome. Cell Mol Life Sci, 2011, 68: 2013-2037.
doi: 10.1007/s00018-011-0673-y pmid: 21611891
[13] 张艺能, 周玉萍, 陈琼华, 等. 拟南芥开花时间调控的分子基础. 植物学报, 2014, 49: 469-482.
doi: 10.3724/SP.J.1259.2014.00469
Zhang Y N, Zhou Y P, Chen Q H, et al. Molecular basis of flowering time regulation in Arabidopsis. Chin Bull Bot, 2014, 49: 469-482 (in Chinese with English abstract).
[14] Ikezaki M, Kojima M, Sakakibara H, et al. Genetic networks regulated by ASYMMETRIC LEAVES1 (AS1) and AS2 in leaf development in Arabidopsis thaliana: KNOX genes control five morphological events. Plant J, 2010, 61: 70-82.
doi: 10.1111/tpj.2009.61.issue-1
[15] Wang H F, Han X F, Fu X F, et al. Overexpression of TaLBD16-4D alters plant architecture and heading date in transgenic wheat. Front Plant Sci, 2022, 13: 911993.
doi: 10.3389/fpls.2022.911993
[16] Wang X F, Zhang S Z, Su L, et al. A genome-wide analysis of the LBD (LATERAL ORGAN BOUNDARIES Domain) gene family in Malus domestica with a functional characterization of MdLBD11. PLoS One, 2013, 8: e57044.
doi: 10.1371/journal.pone.0057044
[17] Gao J S, Niu M X, Lei Y, et al. PomiR164 targets PoLBD25 to promote early flowering in tree peony. Hortic Plant J, Published online [2025-09-06], https://www.sciencedirect.com/science/article/pii/S2468014125001943.
[18] 喻树迅. 我国棉花生产现状与发展趋势. 中国工程科学, 2013, 15(4): 9-13.
Yu S X. Present situation and development trend of cotton production in China. Strategic Study CAE, 2013, 15(4): 9-13 (in Chinese with English abstract).
[19] Li L B, Zhang C, Huang J Q, et al. Genomic analyses reveal the genetic basis of early maturity and identification of loci and candidate genes in upland cotton (Gossypium hirsutum L.). Plant Biotechnol J, 2021, 19: 109-123.
doi: 10.1111/pbi.v19.1
[20] Chang L J, Mei G F, Hu Y, et al. LMI1-like and KNOX1 genes coordinately regulate plant leaf development in dicotyledons. Plant Mol Biol, 2019, 99: 449-460.
doi: 10.1007/s11103-019-00829-7
[21] 崔静. 棉花GbSTKGhLBD100基因抗黄萎病功能研究. 河北农业大学硕士学位论文, 河北保定, 2020.
Cui J. Functional Analysis of GbSTK and GhLBD100 from Cotton in Verticillium wilt resistance. MS Thesis of Hebei Agricultural University, Baoding, Hebei, China, 2020 (in Chinese with English abstract).
[22] 刘醒醒. 陆地棉GhLBD18-GhATG18a调控体细胞胚胎发生细胞自噬的机制研究. 郑州大学硕士学位论文, 河南郑州, 2023.
Liu X X. Mechanism Analysis on GhLBD18-GhATG18a Regulating of Autophagy during Gossypium hirsutum L. Somatic Embryogenesis. MS Thesis of Zhengzhou University, Zhengzhou, Henan, China, 2023 (in Chinese with English abstract).
[23] 王晔. 亚洲棉LBDs同源基因的克隆及应用. 华中农业大学硕士学位论文, 湖北武汉, 2015.
Wang Y. Molecular Cloning and Application of LBDs from Cotton (Gossypium arboreum L.). MS Thesis of Huazhong Agricultural University, Wuhan, Hubei, China, 2015 (in Chinese with English abstract).
[24] Yu J, Jung S, Cheng C H, et al. CottonGen: the community database for cotton genomics, genetics, and breeding research. Plants (Basel), 2021, 10: 2805.
[25] Goodstein D M, Shu S Q, Howson R, et al. Phytozome: a comparative platform for green plant genomics. Nucleic Acids Res, 2012, 40: D1178-D1186.
doi: 10.1093/nar/gkr944
[26] Yates A D, Allen J, Amode R M, et al. Ensembl Genomes 2022: an expanding genome resource for non-vertebrates. Nucleic Acids Res, 2022, 50: D996-D1003.
doi: 10.1093/nar/gkab1007
[27] Finn R D, Mistry J, Schuster-Böckler B, et al. Pfam: clans, web tools and services. Nucleic Acids Res, 2006, 34: D247-D251.
doi: 10.1093/nar/gkj149 pmid: 16381856
[28] Chen C J, Chen H, Zhang Y, et al. TBtools: an integrative toolkit developed for interactive analyses of big biological data. Mol Plant, 2020, 13: 1194-1202.
doi: S1674-2052(20)30187-8 pmid: 32585190
[29] Marchler-Bauer A, Derbyshire M K, Gonzales N R, et al. CDD: NCBI’s conserved domain database. Nucleic Acids Res, 2015, 43: D222-D226.
[30] Kumar S, Stecher G, Suleski M, et al. MEGA12: molecular evolutionary genetic analysis version 12 for adaptive and green computing. Mol Biol Evol, 2024, 41: msae263.
doi: 10.1093/molbev/msae263
[31] Ma Z Y, He S P, Wang X F, et al. Resequencing a core collection of upland cotton identifies genomic variation and loci influencing fiber quality and yield. Nat Genet, 2018, 50: 803-813.
doi: 10.1038/s41588-018-0119-7 pmid: 29736016
[32] Zhang T Z, Hu Y, Jiang W K, et al. Sequencing of allotetraploid cotton (Gossypium hirsutum L. acc. TM-1) provides a resource for fiber improvement. Nat Biotechnol, 2015, 33: 531-537.
doi: 10.1038/nbt.3207
[33] Zhao L J, Lyu Y J, Chen W, et al. Genome-wide identification and analyses of the AHL gene family in cotton (Gossypium). BMC Genom, 2020, 21: 69.
doi: 10.1186/s12864-019-6406-6
[34] Rao X Y, Huang X L, Zhou Z C, et al. An improvement of the 2ˆ (-delta delta CT) method for quantitative real-time polymerase chain reaction data analysis. Biostat Bioinform Biomath, 2013, 3: 71-85.
[35] Guo X F, Yang J N, Li D D, et al. Genome-wide association study reveals novel SNP loci and candidate genes linked to flowering time in upland cotton. Theor Appl Genet, 2025, 138: 214.
doi: 10.1007/s00122-025-05011-w pmid: 40824397
[36] Danecek P, Auton A, Abecasis G, et al. The variant call format and VCFtools. Bioinformatics, 2011, 27: 2156-2158.
doi: 10.1093/bioinformatics/btr330 pmid: 21653522
[37] 程帅帅. 高分辨率的时间动态转录组图谱鉴定棉花开花关键基因及功能验证. 西北农林科技大学博士学位论文, 陕西杨凌, 2020.
Cheng S S. Identification and Functional Analysis of Key Genes in Cotton Flowering by High-Resolution Temporal Dynamic Transcriptome Landscape. PhD Dissertation of Northwest A&F University, Yangling, Shaanxi, China, 2020 (in Chinese with English abstract).
[38] Wang C X, Liu J J, Xie X Y, et al. GhAP1-D3 positively regulates flowering time and early maturity with no yield and fiber quality penalties in upland cotton. J Integr Plant Biol, 2023, 65: 985-1002.
doi: 10.1111/jipb.13409
[39] Zhang X H, Ren Z Y, Hu G H, et al. Functional divergence of GhAP1.1 and GhFUL2 associated with flowering regulation in upland cotton (Gossypium hirsutum L.). J Plant Physiol, 2022, 275: 153757.
doi: 10.1016/j.jplph.2022.153757
[40] Zhang X H, Wei J H, Fan S L, et al. Functional characterization of GhSOC1 and GhMADS42 homologs from upland cotton (Gossypium hirsutum L.). Plant Sci, 2016, 242: 178-186.
doi: 10.1016/j.plantsci.2015.05.001
[41] Cheng S S, Chen P Y, Su Z Z, et al. High-resolution temporal dynamic transcriptome landscape reveals a GhCAL-mediated flowering regulatory pathway in cotton (Gossypium hirsutum L.). Plant Biotechnol J, 2021, 19: 153-166.
doi: 10.1111/pbi.v19.1
[42] Li J, Fan S L, Song M Z, et al. Cloning and characterization of a FLO/LFY ortholog in Gossypium hirsutum L. Plant Cell Rep, 2013, 32: 1675-1686.
doi: 10.1007/s00299-013-1479-1
[43] Yu J W, Xie Q W, Li C, et al. Comprehensive characterization and gene expression patterns of LBD gene family in Gossypium. Planta, 2020, 251: 81.
doi: 10.1007/s00425-020-03364-8
[44] Liu W C, Guo X Y, Zheng T C, et al. Genome-wide identification and characterization of the lateral organ boundaries domain (LBD) gene family in nine Rosaceae species and expression pattern in Prunus mume. Ornament Plant Res, 2024, 4: 1-14.
[45] Sun S, Yi J J, Gu P L, et al. Comprehensive characterization and functional analysis of the lateral organ boundaries domain gene family in rice: evolution, expression, and stress response. Int J Mol Sci, 2025, 26: 3948.
doi: 10.3390/ijms26093948
[46] Hao L D, Li S F, Dai J, et al. Characterization and expression profiles of the ZmLBD gene family in Zea mays. Mol Biol Rep, 2024, 51: 554.
doi: 10.1007/s11033-024-09483-9
[47] Guo B J, Wang J, Lin S, et al. A genome-wide analysis of the ASYMMETRIC LEAVES2/LATERAL ORGAN BOUNDARIES (AS2/LOB) gene family in barley (Hordeum vulgare L.). J Zhejiang Univ Sci B, 2016, 17: 763-774.
doi: 10.1631/jzus.B1500277
[48] Xu J, Hu P, Tao Y, et al. Genome-wide identification and characterization of the Lateral Organ Boundaries Domain (LBD) gene family in polyploid wheat and related species. PeerJ, 2021, 9: e11811.
doi: 10.7717/peerj.11811
[49] Hou J, Liu M, Yang K, et al. Genetic variation for adaptive evolution in response to changed environments in plants. J Integr Plant Biol, 2025, 67: 2265-2293.
doi: 10.1111/jipb.13961
[50] Rast M I, Simon R. Arabidopsis JAGGED LATERAL ORGANS acts with ASYMMETRIC LEAVES2 to coordinate KNOX and PIN expression in shoot and root meristems. Plant Cell, 2012, 24: 2917-2933.
doi: 10.1105/tpc.112.099978
[51] Lin W C, Shuai B, Springer P S. The Arabidopsis LATERAL ORGAN BOUNDARIES-domain gene ASYMMETRIC LEAVES2 functions in the repression of KNOX gene expression and in adaxial-abaxial patterning. Plant Cell, 2003, 15: 2241-2252.
[52] Guo M J, Thomas J, Collins G, et al. Direct repression of KNOX loci by the ASYMMETRIC LEAVES1 complex of Arabidopsis. Plant Cell, 2008, 20: 48-58.
doi: 10.1105/tpc.107.056127
[53] 徐爱武, 李翼然, 尚斌. 我国植棉区棉花种植现状及其发展建议. 中国棉花加工, 2022(5): 28-30.
Xu A W, Li Y R, Shang B. Present situation and development suggestions of cotton planting in cotton planting areas in China. China Cotton Proc, 2022(5): 28-30 (in Chinese).
[54] 王彩香, 袁文敏, 刘娟娟, 等. 西北内陆早熟陆地棉品种的综合评价及育种演化. 中国农业科学, 2023, 56: 1-21.
doi: 10.3864/j.issn.0578-1752.2023.01.001
Wang C X, Yuan W M, Liu J J, et al. Comprehensive evaluation and breeding evolution of early maturing upland cotton varieties in the northwest inland of China. Sci Agric Sin, 2023, 56: 1-21 (in Chinese with English abstract).
doi: 10.3864/j.issn.0578-1752.2023.01.001
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