Zhang Qing, Hu Weichang, Zhu Fan, Wang Liming, Yu Qingyi, Ming Ray, Zhang Jisen
Center for Genomics and Biotechnology, Haixia Institute of Science and Technology (HIST), Fujian Agriculture and Forestry University, Fuzhou, 350002, China.
Department of Plant Biology, University of Illinois at Urbana-Champaign, Urbana, IL, 61801, USA.
BMC Genomics. 2016 Feb 1;17:88. doi: 10.1186/s12864-016-2419-6.
Sugarcane is an economically important crop contributing to about 80% of the world sugar production. Increasing efforts in molecular biological studies have been performed for improving the sugar yield and other relevant important agronomic traits. However, due to sugarcane's complicated genomes, it is still challenging to study the genetic basis of traits, such as sucrose accumulation. Sucrose transporters (SUTs) are critical for both phloem loading in source tissue and sucrose uptaking in sink tissue, and are considered to be the control points for regulating sucrose storage. However, no genomic study for sugarcane sucrose transporter (SsSUT) families has been reported up to date.
By using comparative genomics and bacterial artificial chromosomes (BACs), six SUT genes were identified and characterized in S. spontaenum. Phylogenetic analyses revealed that the two pairs SsSUTs (SsSUT1/SsSUT3 and SsSUT5/SsSUT6) could be clustered together into two separate monocot specific SUT groups, while SsSUT2 and SsSUT4 were separated into the other two groups, with members from both dicot and monocot species. Gene structure comparison demonstrated that the number and position of exons/introns in SUTs were highly conserved among the close orthologs; in contrast, there were variations among the paralogous SUTs in Sacchuarm. Though with the high polyploidy level, gene allelic haplotype comparative analysis showed that the examined four SsSUT members exhibited conservations of gene structures and amino acid sequences among the allelic haplotypes accompanied by variations of intron sizes. Gene expression analyses were performed for tissues from seedlings under drought stress and mature plants of three Saccharum species (S.officinarnum, S.spotaneum and S.robustum). Both SUT1 and SUT4 expressed abundantly at different conditions. SUT2 had similar expression level in all of the examined tissues, but SUT3 was undetectable. Both of SUT5 and SUT6 had lower expression level than other gene member, and expressed stronger in source leaves and are likely to play roles in phloem loading. In the seeding plant leave under water stress, four genes SUT1, SUT2, SUT4 and SUT5 were detectable. In these detectable genes, SUT1 and SUT4 were down regulated, while, SUT2 and SUT5 were up regulated.
In this study, we presented the first comprehensive genomic study for a whole gene family, the SUT family, in Saccharum. We speculated that there were six SUT members in the S. spotaneum genome. Out of the six members, SsSUTs, SsSUT5 and SsSUT6 were recent duplication genes accompanied by rapid evolution, while, SsSUT2 and SsSUT4 were the ancient members in the families. Despite the high polypoidy genome, functional redundancy may not exist among the SUTs allelic haplotypes supported by the evidence of strong purifying selection of the gene allele. SUT3 could be a low active member in the family because it is undetectable in our study, but it might not be a pseudogene because it harbored integrated gene structure. SUT1 and SUT4 were the main members for the sucrose transporter, while, these SUTs had sub-functional divergence in response to sucrose accumulation and plant development in Saccharum.
甘蔗是一种重要的经济作物,占世界食糖产量的约80%。为提高甘蔗的糖分产量和其他相关重要农艺性状,人们在分子生物学研究方面付出了更多努力。然而,由于甘蔗基因组复杂,研究蔗糖积累等性状的遗传基础仍具有挑战性。蔗糖转运蛋白(SUTs)对于源组织中的韧皮部装载和库组织中的蔗糖吸收都至关重要,被认为是调节蔗糖储存的控制点。然而,迄今为止尚未见关于甘蔗蔗糖转运蛋白(SsSUT)家族的基因组研究报道。
通过比较基因组学和细菌人工染色体(BACs),在割手密中鉴定并表征了6个SUT基因。系统发育分析表明,两对SsSUTs(SsSUT1/SsSUT3和SsSUT5/SsSUT6)可聚在一起形成两个单独的单子叶植物特异的SUT组,而SsSUT2和SsSUT4则被分到另外两组,其中包含双子叶和单子叶物种的成员。基因结构比较表明,直系同源SUTs中外显子/内含子的数量和位置在亲缘关系较近的物种中高度保守;相反,甘蔗中旁系同源SUTs之间存在差异。尽管多倍体水平较高,但基因等位单倍型比较分析表明,所检测的4个SsSUT成员在等位单倍型之间表现出基因结构和氨基酸序列的保守性,同时内含子大小存在差异。对干旱胁迫下幼苗和3个甘蔗品种(热带种、割手密和大茎野生种)成熟植株的组织进行了基因表达分析。SUT1和SUT4在不同条件下均大量表达。SUT2在所有检测组织中的表达水平相似,但未检测到SUT3的表达。SUT5和SUT6的表达水平均低于其他基因成员,且在源叶中表达较强,可能在韧皮部装载中发挥作用。在水分胁迫下的幼苗叶片中,可检测到SUT1、SUT2、SUT4和SUT5这4个基因。在这些可检测到的基因中,SUT1和SUT4表达下调,而SUT2和SUT5表达上调。
在本研究中,我们首次对甘蔗中的一个完整基因家族——SUT家族进行了全面的基因组研究。我们推测割手密基因组中有6个SUT成员。在这6个成员中,SsSUT1、SsSUT5和SsSUT6是近期的重复基因,伴随着快速进化,而SsSUT2和SsSUT4是该家族中的古老成员。尽管基因组多倍体水平较高,但基于基因等位基因强烈纯化选择的证据,SUT等位单倍型之间可能不存在功能冗余。SUT3可能是该家族中一个低活性成员,因为在我们的研究中未检测到其表达,但它可能不是假基因,因为它具有完整的基因结构。SUT1和SUT4是蔗糖转运蛋白的主要成员,而这些SUTs在甘蔗中对蔗糖积累和植物发育的响应中存在亚功能分化。