拟南芥AtSUC2和AtSUC4对理化逆境及外源ABA的响应
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摘要
蔗糖作为高等植物中光合同化物运输的主要形式,其运输方向和分配速率影响植物的生长发育,甚至决定作物的产量。细胞膜上的蔗糖转运蛋白(sucrose transporters, SUCs或SUTs)承担着蔗糖从源细胞的外运、韧皮部的装载和卸载,以及向库细胞装入的功能,从而影响蔗糖的运输方向、速率和分配,其基因表达受多种因子的调控,如内源激素、生物逆境和理化逆境等。因此,深入研究SUCs基因表达与环境因子和激素的关系,是阐明环境胁迫下植物光合产物运输和分配调节机制的关键。
蔗糖转运蛋白由多基因家族编码的。在模式植物拟南芥中,蔗糖转运蛋白基因家族有9个成员,分属于3个亚族：SUT1、SUT2和SUT4。基因敲除研究表明,一些成员之间在功能上存在互补作用,有的基因敲除突变体只在特定条件下发生表现型变化。这暗示着不同家族成员的表达调节机制存在差异,不同成员的基因表达对环境胁迫的响应不同。脱落酸(ABA)是植物体内的重要激素之一,是植物逆境响应的关键激素,有胁迫激素之称,参与许多植物逆境反应的调节,包括光合产物运输和分配的调节。但是,目前少有SUCs基因表达与ABA关系的报道。
本研究以拟南芥为材料,利用生物信息学和实时定量PCR (real-time quantitative reverse transcription PCR, qRT-PCR)技术鉴定出高盐、渗透、低温及干旱胁迫和ABA处理响应的拟南芥蔗糖转运蛋白基因：高亲和／低转运的AtSUC2和低亲和／高转运的AtSUC4。观察理化逆境和ABA处理时,AtSUC2和AtSUC4纯合突变体植株的生长发育变化、蔗糖含量变化、ABA响应基因及ABF相关基因表达变化,从而探讨AtSUC2和AtSUC4在拟南芥逆境响应中的作用、AtSUC2和AtSUC4基因表达与其他蔗糖转运蛋白基因表达之间的关系,以及ABA参与调节SUCs基因表达的可能机制。主要结果如下：
1.通过分析基因芯片数据发现：在高盐、渗透、低温和干旱处理期间,AtSUC2和AtSUC4表达量主要表现为显著上升,说明AtSUC2和AtSUC4受理化逆境的正向调节；AtSUCl和AtSUC5在4种胁迫处理期间,表达量均主要表现为显著下降,表明AtSUC1和AtSUC5受理化逆境的负向调节；AtSUC3表达量仅在处理的个别时间发生明显变化；在胁迫处理期间,AtSUC6、AtSUC1、AtSUC8和AtSUC9表达量几乎不发生变化,说明它们在4种理化逆境下可能发挥次要作用。因此,AtSUC2和AtSUC4可能是拟南芥蔗糖转运蛋白基因家族中对理化逆境正向响应的关键基因。
2.利用qRT-PCR技术分析在高盐、渗透、低温和干旱胁迫下拟南芥野生型(wild-type, WT)植株的AtSUC2和AtSUC4基因表达量变化,发现4种胁迫均诱导AtSUC2和AtSUC4基因的表达,表达量均显著高于对照。在处理期间,AtSUC2和AtSUC4表达量先增加,达到最大值后再下降。
3.利用三引物PCR法和RT-PCR (performing reverse transcription, PCRRT-PCR)法均筛选Atsuc2-1、Atsuc2-3、Atsuc3、Atsuc4-1和Atsuc4-2的纯合突变体,作为后续研究的实验材料。
4.在正常条件下,突变体与WT相比,在萌发和生长方面存在显著差异,表现在萌发率降低、叶片数和叶面积减少,主根长度变短。在高盐、渗透和低温胁迫下,突变体与WT在萌发率方面的差异更加明显。但是,随着胁迫时间的延长,突变体萌发率与WT的差异逐渐变小。这些结果表明,理化逆境只是延缓了突变体和WT种子的萌发,并不抑制种子的最终萌发率。在幼苗期,随着胁迫程度的加深,突变体与WT幼苗生长的差异更加显著,这表明AtSUC2和AtSUC4基因的缺失增大植株对理化逆境的敏感性。
5.在正常条件下,AtSUC2和AtSUC4突变体叶片中的蔗糖含量相近,均显著高于WT植株,根中的蔗糖含量相近,均显著低于WT植株。在高盐、渗透和低温胁迫下,WT叶片和根中蔗糖含量与正常条件相比均明显增加；突变体叶片中的蔗糖含量均显著高于WT,根中的蔗糖含量均显著低于WT,随着胁迫程度的增大,突变体与WT相比,叶片和根中蔗糖含量的差异均变为极显著。这表明逆境抑制了蔗糖从源叶向根的运输和分配,并且AtSUC2和AtSUC4在这个过程中起关键作用。
6.观察AtSUC3突变体中AtSUC2和AtSUC4基因在高盐、渗透、低温和干旱胁迫下的表达量变化,结果表明AtSUC3基因缺失降低了AtSUC2和AtSUC4表达量的增加,说明AtSUC3可能参与了AtSUC2和AtSUC4基因表达的调节。
7.利用芯片数据和qRT-PCR技术研究发现外源ABA处理诱导AtSUC2和AtSUC4基因的高表达。进一步的研究发现AtSUC2和AtSUC4基因缺失提高了在种子萌发期和幼苗期植株对ABA的敏感性,例如种子萌发率降低、叶片数和叶面积减少以及主根长度变短；并且外源ABA处理引起叶片中蔗糖的大量积累,根中蔗糖含量显著下降,抑制蔗糖从源向库的运输。此外,AtSUC2和AtSUC4基因突变抑制一些ABA响应基因、ABFs上游及下游基因的表达。这些结果说明AtSUC2和AtSUC4参与了ABA信号传导的调节。
综上所述,AtSUC2和AtSUC4是拟南芥蔗糖转运蛋白基因家族中响应高盐、渗透、低温、干旱胁迫和外源ABA处理诱导的关键基因。它们的缺失导致源叶片蔗糖的积累、根中蔗糖含量的降低,延迟种子的萌发,抑制幼苗的生长。同时,AtSUC2和AtSUC4缺失还抑制理化逆境和ABA处理诱导的基因表达,以及ABF上下游基因的表达。这些结果表明AtSUC2和AtSUC4是植物适应理化逆境的主要调控因子,并且参与ABA依赖的信号转导途径。此外,AtSUC3基因的缺失突变降低理化逆境和外源ABA处理诱导的AtSUC2和AtSUC4表达量的增加。据推测,AtSUC3是一种蔗糖感受器,在感知细胞蔗糖浓度变化后,调节其它蔗糖转运蛋白的活性。本实验的结果也支持这种观点。
Sucrose is the main transport form of carbohydrate in higher plants. The transport direction and partioning efficiency of sucrose influence the plant growth and even are an important factor limiting grain yield. Sucrose transporters (SUCs or SUTs) in plasmalemma as important carriers are primarily responsible for exporting sucrose from source cells, loading and unloading of phloem, and importing sucrose to sink cells, thereby SUCs impact on transport direction, distributing efficiency and distribution of sucrose; and the expression of SUCs are greatly affected by various factors such as endogenous hormone, biotic stress and physicochemical stress. Thus, the further study on the relationship among expression of SUCs gene, environmental factor and hormone is the key to elucidate regulatory mechanism of photoassimilate transport and partioning under environmental stress.
Sucrose transporters are encoded by a multi-gene family, which is composed of nine AtSUCs in model plant Arabidopsis. They have been identified and can be classified into three distinct subfamilies:SUT1-clade, SUT2-clade and SUT4-clade. The results of gene knockout indicate that complementary effects are detected between some family members and some knockout mutants lead to phenotype variation only under some certain conditions, suggesting that there are differences in the regulatory mechanism of SUCs expression, thus the gene expression of different members also varies in response to environmental stress. ABA is one of important plant hormones and a critical hormone in response to stress, also called stress hormone. It plays a crucial role in plant stress tolerance, including photoassimilate transport and partioning under environmental stress. However, the further study on relationship between SUCs and ABA has not been reported nearly to date.
In this research,Arabidopsis was used as material, through microarray data and real-time quantitative reverse transcription PCR(qRT-PCR) analyses, we found that the high affinity/low capacity AtSUC2gene and low affinity/high capacity AtSUC4gene were key genes in response to salt, osmotic, drought, low temperature and exogenous ABA in Arabidopsis. The physicochemical stresses and ABA treatments induced the changes of growth, sucrose contents, ABA responsive genes and ABF-downstream and upstream genes in AtSUC2and AtSUC4homozygous mutants. Furthermore, we further discussed the effect and mechanism of AtSUC2and AtSUC4in response to physicochemical stress. We have also explored interaction of gene expression among AtSUC2, AtSUC4and other AtSUCs, and possible mechanism that ABA is involved in regulating the expression of AtSUCs. The main results are as follows:
1. By the analysis of microarray data, we found that the expression of AtSUC2and AtSUC4gene almost obviously increased in response to salt, osmotic, drought, low temperature, revealing that AtSUC2and AtSUC4may be positive regulators under physicochemical stresses. The expression of AtSUC1and AtSUC5almost markedly decreased in response to4stresses, indicating that AtSUC1and AtSUC5are possible negative regulators under physicochemical stresses. In addition, the expression of AtSUC3was only induced by the stresses at specific treatment times, whereas hardly significant changes in expression were seen for AtSUC6, AtSUC7, AtSUC8and AtSUC9for any stress treatments, suggesting that these genes might have a minor role in plant stress tolerance. Thus, AtSUC2and AtSUC4may be the key genes that positively respond to physicochemical stress.
2. We further investigated the expression of AtSUC2and AtSUC4by qRT-PCR in WT under salt, osmotic, drought and cold stresses, and found that the expression of two genes enhanced significantly in WT under physicochemical stresses. With the treatment time increasing, the expression of two genes increased first and reached the maximum, and then decreased; and all expression under physicochemical stresses were higher than control. Thses results indicated that4physicochemical stresses induced the high expression of AtSUC2and AtSUC4.
3. The homozygous mutants of Atsuc2-1, Atsuc2-3, Atsuc3, Atsuc4-1and Atsuc4-2were identified precisely by three primers PCR and reverse transcription PCR (RT-PCR), and5mutants were used as experimental materials of our subsequent investigations.
4. In the control condition, there was obviously different performance between mutants and WT, for example, lower germination, fewer leaves, shorter roots, and smaller leaf area. Under salt, osmotic and cold stresses, the difference of germination between WT and mutants was more obvious. With the increase of treatment time, the difference of germination reduced gradually. Physicochemical stresses only delayed but not inhibited the germination of seed, thus3stresses had no effect on the final germination percentage of seeds. During seedling stage, these phenotypic differences became more significant with the increase of stress strength, suggesting that the disruption of the AtSUC2and AtSUC4genes led to hypersensitivity to physicochemical stresses.
5. In the control condition, we found sucrose contents in the mutant shoots closed to each other and were higher than in WT shoots, whereas sucrose contents for the mutant roots were similar and lower than for WT roots. Exposure to salt, osmotic and cold stresses resulted much higher sucrose contents in shoots and roots of WT and mutants compared with sucrose contents in the control condition, much higher sucrose contents in mutant shoots than in WT shoots, and much lower sucrose contents in mutant roots than in WT roots, and the difference was statistically significant. The results indicate that physicochemical stresses inhibit the transport and partioning of sucrose from source leaves to roots, and AtSUC2and AtSUC4play a critical role in the process.
6. By investigating AtSUC2and AtSUC4expression in Atsuc3under normal and physicochemical stress conditions, we found that disruption of AtSUC3weakened the increase in AtSUC2and AtSUC4expression induced by salt, osmotic, drought and cold stresses, indicating that AtSUC3may be involved in modulating the expression of AtSUC2and AtSUC4under physicochemical stress.
7. We investigated the expression of AtSUC2and AtSUC4by microarray data and qRT-PCR in WT under exogenous ABA treatment, the results showed that exogenous ABA induced the high expression of AtSUC2and AtSUC4. We found that loss-of-function mutants of AtSUC2and AtSUC4showed hypersensitivity to exogenous ABA during seed germination and seedling growth, such as lower germination, fewer leaves, shorter roots, and smaller leaf area. Exogenous ABA treatments induced higher sucrose content in shoots and lower sucrose content in roots of these mutants compared with WT, revealing that the disruption of the AtSUC2and AtSUC4inhibit the transport of sucrose from source leaves to roots. In addition, disruption of the AtSUC2and AtSUC4also inhibited the ABA-induced expression of many stressES and ABA responsive genes, especially ABFs and ABF-downstream and upstream genes. The results show that AtSUC2and AtSUC4are involved in ABA signal transduction.
In sum, AtSUC2and AtSUC4are key genes in AtSUC gene family in response to salt, osmotic, drought, cold stresses and exogenous ABA. The loss-of-function mutants of AtSUC2and AtSUC4resulted in accumulation of sucrose in source leaves, decreasing of sucrose contents in roots, the delay of seed germination and inhibition of seeding growth. Furthermore, disruption of the AtSUC2and AtSUC4also inhibited the ABA responsive genes and ABF-downstream and upstream genes. These findings confirmed that AtSUC2and AtSUC4are important regulators in plant physicochemical stress tolerance and operate an ABA-dependent signalling pathway. In addition, disruption of AtSUC3weakened the increase in AtSUC2and AtSUC4expression induced by physicochemical stresses and exogenous ABA. Presumably, AtSUC3is a sucrose sensor, sensing the changes of sucrose concentration in cell and regulating the activity of other AtSUCs; the results in this research also support this opinion.
蔗糖转运蛋白由多基因家族编码的。在模式植物拟南芥中,蔗糖转运蛋白基因家族有9个成员,分属于3个亚族：SUT1、SUT2和SUT4。基因敲除研究表明,一些成员之间在功能上存在互补作用,有的基因敲除突变体只在特定条件下发生表现型变化。这暗示着不同家族成员的表达调节机制存在差异,不同成员的基因表达对环境胁迫的响应不同。脱落酸(ABA)是植物体内的重要激素之一,是植物逆境响应的关键激素,有胁迫激素之称,参与许多植物逆境反应的调节,包括光合产物运输和分配的调节。但是,目前少有SUCs基因表达与ABA关系的报道。
本研究以拟南芥为材料,利用生物信息学和实时定量PCR (real-time quantitative reverse transcription PCR, qRT-PCR)技术鉴定出高盐、渗透、低温及干旱胁迫和ABA处理响应的拟南芥蔗糖转运蛋白基因：高亲和／低转运的AtSUC2和低亲和／高转运的AtSUC4。观察理化逆境和ABA处理时,AtSUC2和AtSUC4纯合突变体植株的生长发育变化、蔗糖含量变化、ABA响应基因及ABF相关基因表达变化,从而探讨AtSUC2和AtSUC4在拟南芥逆境响应中的作用、AtSUC2和AtSUC4基因表达与其他蔗糖转运蛋白基因表达之间的关系,以及ABA参与调节SUCs基因表达的可能机制。主要结果如下：
1.通过分析基因芯片数据发现：在高盐、渗透、低温和干旱处理期间,AtSUC2和AtSUC4表达量主要表现为显著上升,说明AtSUC2和AtSUC4受理化逆境的正向调节；AtSUCl和AtSUC5在4种胁迫处理期间,表达量均主要表现为显著下降,表明AtSUC1和AtSUC5受理化逆境的负向调节；AtSUC3表达量仅在处理的个别时间发生明显变化；在胁迫处理期间,AtSUC6、AtSUC1、AtSUC8和AtSUC9表达量几乎不发生变化,说明它们在4种理化逆境下可能发挥次要作用。因此,AtSUC2和AtSUC4可能是拟南芥蔗糖转运蛋白基因家族中对理化逆境正向响应的关键基因。
2.利用qRT-PCR技术分析在高盐、渗透、低温和干旱胁迫下拟南芥野生型(wild-type, WT)植株的AtSUC2和AtSUC4基因表达量变化,发现4种胁迫均诱导AtSUC2和AtSUC4基因的表达,表达量均显著高于对照。在处理期间,AtSUC2和AtSUC4表达量先增加,达到最大值后再下降。
3.利用三引物PCR法和RT-PCR (performing reverse transcription, PCRRT-PCR)法均筛选Atsuc2-1、Atsuc2-3、Atsuc3、Atsuc4-1和Atsuc4-2的纯合突变体,作为后续研究的实验材料。
4.在正常条件下,突变体与WT相比,在萌发和生长方面存在显著差异,表现在萌发率降低、叶片数和叶面积减少,主根长度变短。在高盐、渗透和低温胁迫下,突变体与WT在萌发率方面的差异更加明显。但是,随着胁迫时间的延长,突变体萌发率与WT的差异逐渐变小。这些结果表明,理化逆境只是延缓了突变体和WT种子的萌发,并不抑制种子的最终萌发率。在幼苗期,随着胁迫程度的加深,突变体与WT幼苗生长的差异更加显著,这表明AtSUC2和AtSUC4基因的缺失增大植株对理化逆境的敏感性。
5.在正常条件下,AtSUC2和AtSUC4突变体叶片中的蔗糖含量相近,均显著高于WT植株,根中的蔗糖含量相近,均显著低于WT植株。在高盐、渗透和低温胁迫下,WT叶片和根中蔗糖含量与正常条件相比均明显增加；突变体叶片中的蔗糖含量均显著高于WT,根中的蔗糖含量均显著低于WT,随着胁迫程度的增大,突变体与WT相比,叶片和根中蔗糖含量的差异均变为极显著。这表明逆境抑制了蔗糖从源叶向根的运输和分配,并且AtSUC2和AtSUC4在这个过程中起关键作用。
6.观察AtSUC3突变体中AtSUC2和AtSUC4基因在高盐、渗透、低温和干旱胁迫下的表达量变化,结果表明AtSUC3基因缺失降低了AtSUC2和AtSUC4表达量的增加,说明AtSUC3可能参与了AtSUC2和AtSUC4基因表达的调节。
7.利用芯片数据和qRT-PCR技术研究发现外源ABA处理诱导AtSUC2和AtSUC4基因的高表达。进一步的研究发现AtSUC2和AtSUC4基因缺失提高了在种子萌发期和幼苗期植株对ABA的敏感性,例如种子萌发率降低、叶片数和叶面积减少以及主根长度变短；并且外源ABA处理引起叶片中蔗糖的大量积累,根中蔗糖含量显著下降,抑制蔗糖从源向库的运输。此外,AtSUC2和AtSUC4基因突变抑制一些ABA响应基因、ABFs上游及下游基因的表达。这些结果说明AtSUC2和AtSUC4参与了ABA信号传导的调节。
综上所述,AtSUC2和AtSUC4是拟南芥蔗糖转运蛋白基因家族中响应高盐、渗透、低温、干旱胁迫和外源ABA处理诱导的关键基因。它们的缺失导致源叶片蔗糖的积累、根中蔗糖含量的降低,延迟种子的萌发,抑制幼苗的生长。同时,AtSUC2和AtSUC4缺失还抑制理化逆境和ABA处理诱导的基因表达,以及ABF上下游基因的表达。这些结果表明AtSUC2和AtSUC4是植物适应理化逆境的主要调控因子,并且参与ABA依赖的信号转导途径。此外,AtSUC3基因的缺失突变降低理化逆境和外源ABA处理诱导的AtSUC2和AtSUC4表达量的增加。据推测,AtSUC3是一种蔗糖感受器,在感知细胞蔗糖浓度变化后,调节其它蔗糖转运蛋白的活性。本实验的结果也支持这种观点。
Sucrose is the main transport form of carbohydrate in higher plants. The transport direction and partioning efficiency of sucrose influence the plant growth and even are an important factor limiting grain yield. Sucrose transporters (SUCs or SUTs) in plasmalemma as important carriers are primarily responsible for exporting sucrose from source cells, loading and unloading of phloem, and importing sucrose to sink cells, thereby SUCs impact on transport direction, distributing efficiency and distribution of sucrose; and the expression of SUCs are greatly affected by various factors such as endogenous hormone, biotic stress and physicochemical stress. Thus, the further study on the relationship among expression of SUCs gene, environmental factor and hormone is the key to elucidate regulatory mechanism of photoassimilate transport and partioning under environmental stress.
Sucrose transporters are encoded by a multi-gene family, which is composed of nine AtSUCs in model plant Arabidopsis. They have been identified and can be classified into three distinct subfamilies:SUT1-clade, SUT2-clade and SUT4-clade. The results of gene knockout indicate that complementary effects are detected between some family members and some knockout mutants lead to phenotype variation only under some certain conditions, suggesting that there are differences in the regulatory mechanism of SUCs expression, thus the gene expression of different members also varies in response to environmental stress. ABA is one of important plant hormones and a critical hormone in response to stress, also called stress hormone. It plays a crucial role in plant stress tolerance, including photoassimilate transport and partioning under environmental stress. However, the further study on relationship between SUCs and ABA has not been reported nearly to date.
In this research,Arabidopsis was used as material, through microarray data and real-time quantitative reverse transcription PCR(qRT-PCR) analyses, we found that the high affinity/low capacity AtSUC2gene and low affinity/high capacity AtSUC4gene were key genes in response to salt, osmotic, drought, low temperature and exogenous ABA in Arabidopsis. The physicochemical stresses and ABA treatments induced the changes of growth, sucrose contents, ABA responsive genes and ABF-downstream and upstream genes in AtSUC2and AtSUC4homozygous mutants. Furthermore, we further discussed the effect and mechanism of AtSUC2and AtSUC4in response to physicochemical stress. We have also explored interaction of gene expression among AtSUC2, AtSUC4and other AtSUCs, and possible mechanism that ABA is involved in regulating the expression of AtSUCs. The main results are as follows:
1. By the analysis of microarray data, we found that the expression of AtSUC2and AtSUC4gene almost obviously increased in response to salt, osmotic, drought, low temperature, revealing that AtSUC2and AtSUC4may be positive regulators under physicochemical stresses. The expression of AtSUC1and AtSUC5almost markedly decreased in response to4stresses, indicating that AtSUC1and AtSUC5are possible negative regulators under physicochemical stresses. In addition, the expression of AtSUC3was only induced by the stresses at specific treatment times, whereas hardly significant changes in expression were seen for AtSUC6, AtSUC7, AtSUC8and AtSUC9for any stress treatments, suggesting that these genes might have a minor role in plant stress tolerance. Thus, AtSUC2and AtSUC4may be the key genes that positively respond to physicochemical stress.
2. We further investigated the expression of AtSUC2and AtSUC4by qRT-PCR in WT under salt, osmotic, drought and cold stresses, and found that the expression of two genes enhanced significantly in WT under physicochemical stresses. With the treatment time increasing, the expression of two genes increased first and reached the maximum, and then decreased; and all expression under physicochemical stresses were higher than control. Thses results indicated that4physicochemical stresses induced the high expression of AtSUC2and AtSUC4.
3. The homozygous mutants of Atsuc2-1, Atsuc2-3, Atsuc3, Atsuc4-1and Atsuc4-2were identified precisely by three primers PCR and reverse transcription PCR (RT-PCR), and5mutants were used as experimental materials of our subsequent investigations.
4. In the control condition, there was obviously different performance between mutants and WT, for example, lower germination, fewer leaves, shorter roots, and smaller leaf area. Under salt, osmotic and cold stresses, the difference of germination between WT and mutants was more obvious. With the increase of treatment time, the difference of germination reduced gradually. Physicochemical stresses only delayed but not inhibited the germination of seed, thus3stresses had no effect on the final germination percentage of seeds. During seedling stage, these phenotypic differences became more significant with the increase of stress strength, suggesting that the disruption of the AtSUC2and AtSUC4genes led to hypersensitivity to physicochemical stresses.
5. In the control condition, we found sucrose contents in the mutant shoots closed to each other and were higher than in WT shoots, whereas sucrose contents for the mutant roots were similar and lower than for WT roots. Exposure to salt, osmotic and cold stresses resulted much higher sucrose contents in shoots and roots of WT and mutants compared with sucrose contents in the control condition, much higher sucrose contents in mutant shoots than in WT shoots, and much lower sucrose contents in mutant roots than in WT roots, and the difference was statistically significant. The results indicate that physicochemical stresses inhibit the transport and partioning of sucrose from source leaves to roots, and AtSUC2and AtSUC4play a critical role in the process.
6. By investigating AtSUC2and AtSUC4expression in Atsuc3under normal and physicochemical stress conditions, we found that disruption of AtSUC3weakened the increase in AtSUC2and AtSUC4expression induced by salt, osmotic, drought and cold stresses, indicating that AtSUC3may be involved in modulating the expression of AtSUC2and AtSUC4under physicochemical stress.
7. We investigated the expression of AtSUC2and AtSUC4by microarray data and qRT-PCR in WT under exogenous ABA treatment, the results showed that exogenous ABA induced the high expression of AtSUC2and AtSUC4. We found that loss-of-function mutants of AtSUC2and AtSUC4showed hypersensitivity to exogenous ABA during seed germination and seedling growth, such as lower germination, fewer leaves, shorter roots, and smaller leaf area. Exogenous ABA treatments induced higher sucrose content in shoots and lower sucrose content in roots of these mutants compared with WT, revealing that the disruption of the AtSUC2and AtSUC4inhibit the transport of sucrose from source leaves to roots. In addition, disruption of the AtSUC2and AtSUC4also inhibited the ABA-induced expression of many stressES and ABA responsive genes, especially ABFs and ABF-downstream and upstream genes. The results show that AtSUC2and AtSUC4are involved in ABA signal transduction.
In sum, AtSUC2and AtSUC4are key genes in AtSUC gene family in response to salt, osmotic, drought, cold stresses and exogenous ABA. The loss-of-function mutants of AtSUC2and AtSUC4resulted in accumulation of sucrose in source leaves, decreasing of sucrose contents in roots, the delay of seed germination and inhibition of seeding growth. Furthermore, disruption of the AtSUC2and AtSUC4also inhibited the ABA responsive genes and ABF-downstream and upstream genes. These findings confirmed that AtSUC2and AtSUC4are important regulators in plant physicochemical stress tolerance and operate an ABA-dependent signalling pathway. In addition, disruption of AtSUC3weakened the increase in AtSUC2and AtSUC4expression induced by physicochemical stresses and exogenous ABA. Presumably, AtSUC3is a sucrose sensor, sensing the changes of sucrose concentration in cell and regulating the activity of other AtSUCs; the results in this research also support this opinion.
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