彩色棉与白色棉产量和品质差异的分子生理机制研究
摘要
天然彩色棉产量偏低和品质偏差是其在生产利用上需要解决的重要问题之一。天然彩色棉纤维发育过程中,除纤维素合成外,色素代谢的增加是区别白色棉纤维发育的重要特征之一。纤维中色素的合成将导致纤维细胞中碳水化合物的消耗,从而影响到纤维素的合成。为此,本文以底物竞争为中心点,从天然彩色棉与白色棉植株光合特性、纤维碳水化合物和色素含量变化、纤维细胞呼吸代谢和碳水化合物代谢相关酶的功能和基因表达等角度分析了造成产量和品质差异的分子生理机制。主要研究结果如下:
1、天然彩色棉的光合特性与产量和品质的关系
以棕色棉、绿色棉和两个棕色棉杂种:棕杂1(陆陆杂种)和棕杂2(海陆杂种)及白色棉对照为材料,研究了棉株不同发育时期叶片光合色素含量、叶绿体希尔活性、Mg~(2+)-ATPase和Ca~(2+)-ATPase活性、净光合速率和淀粉、蔗糖、果糖和葡萄糖等碳水化合物含量的变化,并对这些光合指标与棉花产量和品质间进行了相关性和通径分析。结果表明,绿色棉的产量最低,仅为白色棉的58%,其次是棕色棉,为白色棉的66%。棕杂1的产量最高,棕杂2介于棕杂和白色棉之间。绿色棉虽然叶片中的光合色素含量、净光合速率和其他光合指标与白色棉相当,甚至更高,但其产量仍然最低,引起绿色棉产量和品质低的主要原冈是营养生长过旺,单株铃数少,单铃重轻:相反,棕色棉产量低和品质较差的重要原因则是由于其光合作用弱,叶片中叶绿体生理活性不强,光合产物少。就两个杂种而言,棕杂1的光合能力要强于棕杂2,而且棕杂1的产量也显著高于棕杂2,但其品质却比棕杂2略差。棉株光合特性与天然彩色棉产量与品质之间的相关性和通径分析结果表明,净光合速率对棉花产量的贡献最大,其次为碳水化合物;而纤维长度则是叶片中的碳水化合物含量对其影响最大,净光合速率次之,这与棉花纤维发育过程中碳水化合物除用于原料支持各种代谢活动外,还起到调节纤维细胞渗透压,促进纤维伸长有关,尤其用于纤维素的合成,因为纤维素含量占总棉纤维的重量95%左右。研究结果表明,通过选育具有高光效的杂交组合提高天然彩色棉纤维的产量和品质是可行的。
2、天然彩色棉纤维碳水化合物与色素含量对纤维长度的影响
对白色棉、棕色棉、绿色棉、棕色棉陆陆杂种和海陆杂种纤维发育过程中碳水化合物、色素含量、纤维素含量和纤维长度动态进行了检测。结果表明,从开花当天至纤维成熟的整个发育过程中,白色棉和天然彩色棉纤维中的蔗糖含量在10 DPA前含量最高,之后持续降低;果糖含量从0 DPA至成熟一直处于减少状态,而葡萄糖含量则在20 DPA,即纤维素合成高峰期达到最高。绿色棉纤维和棕杂1纤维的糖含量相当,均比其他类型棉花高。但绿色棉纤维的产量和品质最低。这是因为其纤维中PAL活性高,导致色素大量合成,碳水化合物的消耗量较大,从而影响到纤维素的合成。而且,其碳水化合物的利用率低(马克隆值低,纤维中碳水化合物滞留量大),也是其纤维素合成量少的重要因素。另外,棕色棉纤维碳水化合物含量最少,因此,纤维素合成的原料少,加之部分碳水化合物用于色素的合成,导致其纤维产量和品质比白色棉低。陆陆杂种纤维中较高的碳水化合物含量可以补偿用于色素合成所消耗的部分碳水化合物含量。可见,天然彩色棉纤维中色素和纤维素的合成在碳水化合物的利用上存在着此消彼长的关系。
3、天然彩色棉纤维细胞氧化还原状态与外源呼吸抑制剂对纤维发育的影响
通过测定白色棉、棕色棉、绿色棉和两个棕色棉杂种纤维细胞发育过程中细胞含水量、pH值、细胞色素c氧化酶活性、NAD~+、NADH、NADP~+、和NADPH的含量及在组培条件下研究了鱼藤酮和硫脲两种呼吸抑制剂对白色棉和绿色棉纤维显色、胚珠鲜重和纤维长度及细胞色素c氧化酶和多酚氧化酶活性的影响。结果表明,绿色棉和棕色棉纤维细胞的显色时间和方式存在明显的差异。棕色棉在30 DPA后以“爆发式”显色;绿色棉则在20 DPA时以“渐进式”逐步使绿色加深,因此,棕色棉和绿色棉纤维中的生理代谢状况是不一样的。白色棉纤维中的水分含量较高,其纤维细胞中的pH值比绿色棉低,但比棕色棉系列棉花高。棕色棉系列纤维细胞中的细胞色素c氧化酶、NAD~+、NADH、NADP~+、和NADPH含量均远远高于白色棉和绿色棉,而且NAD~+或NADP~+含量比NADH或NADPH高,因此,其纤维细胞中的呼吸代谢旺盛;白色棉和绿色棉纤维细胞中的辅酶含量较低,而且两者差异不大。绿色棉在呼吸代谢强度上与白色棉相当,但综合纤维中的色素和纤维素合成两大重要代谢,因此,可以推断绿色棉纤维中大量碳水化合物可能未能被充分利用:相反棕色棉虽然呼吸代谢旺盛,但纤维细胞中的碳水化合物含量少,故造成其纤维素含量比白色棉低,但比绿色棉高;而色素含量比白色棉高,但比绿色棉低。鱼藤酮和硫脲均可抑制白色棉和绿色棉胚珠和纤维细胞的发育,胚珠鲜重减轻,纤维减短,但鱼藤酮的毒害作用更大,而且可使绿色棉纤维显色较对照(绿色棉末用鱼藤酮处理)提早1周;硫脲对胚珠和纤维的发育伤害虽然较小,但可使绿色棉纤维不显色,显然,硫脲对绿色棉纤维的色素代谢具有抑制作用。在鱼藤酮处理下,纤维细胞中的细胞色素c氧化酶的活性迅速降低,而在硫脲各浓度处理下,绿色棉纤维中的多酚氧化酶活性均降低了80%以上。
4、天然彩色棉纤维细胞碳水化合物代谢关键酶活性及基因表达研究
本试验以白色棉、棕色棉、绿色棉和2个棕色棉杂种为材料研究了纤维0-20 DPA发育阶段纤维中酸性和中性转化酶和蔗糖合成酶、蔗糖磷酸合成酶的活性。并且采用RT-PCR和荧光定量RT-PCR的方法对蔗糖合成酶基因的表达量进行了测定。结果表明,棕色棉纤维细胞中的转化酶和蔗糖合成酶最高,绿色棉和白色棉纤维细胞中转化酶和蔗糖合成酶活性均较低相差不大。在2 DPA至20 DPA期间,白色棉和天然彩色棉纤维细胞中的蔗糖合成酶活性高且变化幅度很小,说明纤维细胞发育过程中碳水化合物代谢非常旺盛。白色棉、棕色棉和绿色棉纤维细胞中蔗糖合成酶基因的表达量随纤维发育时期的推进变化不大,但棕色棉纤维细胞中的蔗糖合成酶基因的表达量比白色棉高,绿色棉相对较低。此外,白色棉、棕色棉和绿色棉纤维细胞中的蔗糖合成酶基因的表达量比胚珠和叶片的表达量高。绿色棉纤维细胞中蔗糖合成酶活性和基因表达量低是其碳水化合物利用率低的重要原因。
5、棉花纤维发育特异启动子的克隆及其应用
从棉花基因组DNA中分离到棉花纤维发育特异基因GaRDL 1的启动子,并在其后连接绿色荧光蛋白作为报告基因,采用花粉管通道法转化胚珠。结果表明,在子房注射3天后可在纤维中检测到绿色荧光蛋白,表明该基因启动子在棉花纤维中特异表达。利用反义抑制的方法,在启动子后面连接蔗糖合成酶反义基因,并采用花粉管通道法转化胚珠后,取子房注射3天后的胚珠与纤维的混合物进行蔗糖合成酶活性和碳水化合物含量的测定。结果表明,白色棉、棕色棉和绿色棉纤维中的蔗糖合成酶活性降低,但蔗糖含量因未被蔗糖合成酶分解,因此,蔗糖含量变化不大,同时导致纤维中的葡萄糖和果糖含量降低,从而降低了纤维的碳水化合物利用率。
Low yield and quality of naturally colored cotton fiber is one of the most important problems that is required to be solved in the genetic improvement. During fiber development, besides cellulose synthesis, pigment metabolism is an additional process in naturally colored cotton fiber that differed in white cotton fiber. Pigment synthesis in fiber will consume part of carbohydrate in fiber cells affecting cellulose synthesis for fiber yield and quality. Therefore, in present paper, the molecular and physiological mechanism for the difference of yield and quality between white and naturally colored cotton fibers were analyzed from plant photosynthesis, fiber carbohydrate and pigment content, fiber respiration, and gene function and expression that related with carbohydrate metabolic enzymes aspects according to substrate competition hypothesis. The main results wereshowed as below.
1. Relationship between photosynthetic property and yield and quality of naturally colored cotton fiber
Using brown fiber cotton, green fiber cotton, two brown hybrids, hybrid 1 (the parents were upland cotton) and hybrid 2 (the two parents were upland cotton and sea island cotton), white fiber cotton (control) as plant material, variation of photosynthetic pigment content, chloroplast Hill reaction activity, Mg~(2+)-ATPase and Ca~(2+)-ATPase activity, net photosynthetic rate, leaf starch, sucrose, fructose, and glucose content was determined. Furthermore, correlation and path analysis among yield and quality and these photosynthetic indexes were also analyzed. Result showed that yield ranking of cotton fiber was hybrid 1>white fiber cotton>hybrid 2> brown fiber cotton>green fiber cotton. Yield of green fiber cotton and brown fiber cotton was only 58% and 66% of white fiber cotton, respectively. Although leaf photosynthetic pigment content, net photosynthetic rate and other photosynthetic indexes of green fiber cotton plant was near to white fiber cotton even much higher, its yield was yet the lowest. Lower yield and quality in green fiber cotton was mainly caused by excessively nutritional plant growth, less bolls per plant, and lighter boll mass. However, the reason for low yield and quality of brown fiber cotton was its weak photosynthesis, less physiological activity of leaf chloroplast, and small amount of photosynthetic production. As for the two brown hybrids, photosynthetic ability of hybrid 1 much better than hybrid 2 and yield of hybrid 1 was significantly higher than hybrid 2. However, quality of hybrid 2 was slightly better than hybrid 1. Result of correlation and path analysis among yield and quality and photosynthetic indexes indicated that net photosynthetic rate had the most contribution to yield and the next important was carbohydrate content while had the reverse tendency for fiber length, which was possibly due to its fiber cellular osmotic regulation promoting fiber elongation besides it was used as metabolic substrate. These results indicated that it was feasible for fiber yield and quality improvement through screening for efficiency photosynthesis hybrid combination.
2. Effect of fiber carbohydrate and pigment content on fiber length
Fiber carbohydrate and pigment content, cellulose content, and fiber length dynamic were investigated in white fiber cotton, brown fiber cotton, green fiber cotton, and two brown hybrids during fiber development. Result showed that fiber sucrose content was the highest at 10 DPA and then continuously decreased in white fiber cotton and naturally fiber cotton from the day of anthesis to fiber maturation. Fiber fructose content kept decreasing state from 0 DPA to fiber maturation. However fiber glucose content peaked at 20 DPA, which was consistent with the vigorous stage of the fiber cellulose synthesis both in white and naturally fiber cotton. Fiber carbohydrate content in green fiber cotton and brown hybrid 1 was similar and much higher than other type cotton. However, green fiber cotton had the lowest yield and quality. It was because higher PAL activity leading to large amount of pigment synthesis in green cotton fiber, which would consume much carbohydrate. Thus cellulose synthetic amount reduced. In addition to this reason, low carbohydrate use efficiency was other possibly important reason for low cellulose content because of its low micronaire value, which indicated much carbohydrate was not to be used in cellulose synthesis as fiber maturation. In brown cotton fiber, its lower cellulose content than white cotton fiber was due to low carbohydrate content in fiber and consumption of part carbohydrate for pigment synthesis resulting in substrate reduction. As for hybrid 1, much higher carbohydrate content in fiber could partly make up for the usage of carbohydrate in pigment synthesis. These results demonstrated the substrate ebbing and flowing phenomenon in carbohydrate consumption for pigment and cellulose synthesis.
3. Impact of cellular redox state and exogenous respiratory inhibitors on fiber development
In this section, cellular water content, pH value, cytochrome c oxidase activity, NAD~+, NADH, NADP~+, and NADPH content was monitored as fiber cell development. Furthermore, color display, ovule fresh weight, fiber length, cytochrome c oxidase and polyphenoloxidase activity in green and white cotton fiber were also traced under two respiratory inhibitor treatment, rotenone and thiourea, with in vitro culture condition. Result showed that color display timing and manner was obviously different in green and brown cotton fiber. It displayed as "suddenly" style after 30 DPA in brown cotton fiber while as "increasingly" after 20 DPA in green cotton fiber. Therefore, it could be inferred that their physiological state was different in brown and green cotton fiber. White cotton fiber had much higher water content. However, its pH value was lower than green cotton fiber and higher than brown cotton fiber. Cytochrome c oxidase activity, NAD~+, NADH, NADP~+, and NADPH content in brown serials of cotton fiber was far higher than white and green cotton fiber and NAD~+ or NADP~+ content was much higher than NADH or NADPH demonstrating vigorous respiration in fiber cells. However, coenzyme content in white and green fiber was relatively low and little difference between them. Because green cotton fiber contained two important metabolic systems, which were pigment and cellulose synthesis, it could be inferred lots of carbohydrate might be fully used. Although high activity of respiratory metabolism in brown cotton fiber, cellulose content was higher than green cotton fiber and lower than white cotton fiber while pigment content reversely because of less amount of carbohydrate content in fiber. Ovule and fiber development would be inhibited by both rotenone and thiourea causing ovule fresh weight reduction and fiber shortening. But rotenone was more harmful to ovule and fiber development and color displayed for one week earlier in green cotton fiber. Although thiourea had less effect on ovule and fiber development, it could result in no color display in green cotton fiber indicating pigment metabolism was inhibited by thiourea. Moreover, cytochrome c oxidase activity rapidly decreased in fiber cells after rotenone treatment and polyphenoloxidase activity reduced by more than 80% in green cotton fiber after thiourea treatment.
4. Activity of key enzymes in carbohydrate metabolism in naturally cotton fibers and sucrose synthase gene expression
Carbohydrate metabolic related enzymes, i.e. acidic and neutral invertase, sucrose synthase, and sucrose phosphate synthase, were measured in white, brown, green cotton fiber, and two brown hybrids when fiber was at 0-20 DPA developmental stages. Furthermore, expression amount of sucrose synthase gene was assayed using RT-PCR and real time RT-PCR method. Result showed that activity invertase and sucrose synthase was the highest in brown cotton fiber. These enzymatic activities were similar between green and white cotton fiber. Sucrose synthase activity in white and naturally colored cotton fiber varied little from 2 DPA o 20 DPA revealing high activity of carbohydrate metabolism during cotton fiber development. Furthermore, sucrose synthase gene expression in white, brown, and green cotton fiber altered little during fiber development which was in accordance with enzyme activity variation. However, expression level of sucrose synthase in brown cotton fiber was higher than white cotton fiber. Green cotton fiber had the lowest expression level in fiber cells. As for different organs, fiber cells had much higher expression level than ovule and leaf in white, brown, and green fiber cotton. Low activity of sucrose synthase and gene expression in green cotton fiber was an important reason for its low utilization efficiency of carbohydrate.
5. Molecular cloning of cotton fiber development specific promoter and its application
A cotton fiber development specific promoter of GaRDL 1 gene had been isolated from cotton genomic DNA. The promoter was connected GFP sequentially as report gene and was transformed into ovule by pollen tube pathway. Result showed that GFP could be detected in fiber after 3 days of ovary injection uncovering the specific expression of promoter in fiber. Sucrose synthase gene was reversely ligased after promoter and sucrose synthase activity and carbohydrate content of mixture of ovule and fiber was measured after 3 days of ovary injection using antisense inhibition method. Result showed that sucrose synthase activity reduced in white, brown, and green cotton fiber. However, sucrose content changes little because of its low degradation resulting in low content of glucose and fructose. And thus carbohydrate utilization efficiency was lowered.
1、天然彩色棉的光合特性与产量和品质的关系
以棕色棉、绿色棉和两个棕色棉杂种:棕杂1(陆陆杂种)和棕杂2(海陆杂种)及白色棉对照为材料,研究了棉株不同发育时期叶片光合色素含量、叶绿体希尔活性、Mg~(2+)-ATPase和Ca~(2+)-ATPase活性、净光合速率和淀粉、蔗糖、果糖和葡萄糖等碳水化合物含量的变化,并对这些光合指标与棉花产量和品质间进行了相关性和通径分析。结果表明,绿色棉的产量最低,仅为白色棉的58%,其次是棕色棉,为白色棉的66%。棕杂1的产量最高,棕杂2介于棕杂和白色棉之间。绿色棉虽然叶片中的光合色素含量、净光合速率和其他光合指标与白色棉相当,甚至更高,但其产量仍然最低,引起绿色棉产量和品质低的主要原冈是营养生长过旺,单株铃数少,单铃重轻:相反,棕色棉产量低和品质较差的重要原因则是由于其光合作用弱,叶片中叶绿体生理活性不强,光合产物少。就两个杂种而言,棕杂1的光合能力要强于棕杂2,而且棕杂1的产量也显著高于棕杂2,但其品质却比棕杂2略差。棉株光合特性与天然彩色棉产量与品质之间的相关性和通径分析结果表明,净光合速率对棉花产量的贡献最大,其次为碳水化合物;而纤维长度则是叶片中的碳水化合物含量对其影响最大,净光合速率次之,这与棉花纤维发育过程中碳水化合物除用于原料支持各种代谢活动外,还起到调节纤维细胞渗透压,促进纤维伸长有关,尤其用于纤维素的合成,因为纤维素含量占总棉纤维的重量95%左右。研究结果表明,通过选育具有高光效的杂交组合提高天然彩色棉纤维的产量和品质是可行的。
2、天然彩色棉纤维碳水化合物与色素含量对纤维长度的影响
对白色棉、棕色棉、绿色棉、棕色棉陆陆杂种和海陆杂种纤维发育过程中碳水化合物、色素含量、纤维素含量和纤维长度动态进行了检测。结果表明,从开花当天至纤维成熟的整个发育过程中,白色棉和天然彩色棉纤维中的蔗糖含量在10 DPA前含量最高,之后持续降低;果糖含量从0 DPA至成熟一直处于减少状态,而葡萄糖含量则在20 DPA,即纤维素合成高峰期达到最高。绿色棉纤维和棕杂1纤维的糖含量相当,均比其他类型棉花高。但绿色棉纤维的产量和品质最低。这是因为其纤维中PAL活性高,导致色素大量合成,碳水化合物的消耗量较大,从而影响到纤维素的合成。而且,其碳水化合物的利用率低(马克隆值低,纤维中碳水化合物滞留量大),也是其纤维素合成量少的重要因素。另外,棕色棉纤维碳水化合物含量最少,因此,纤维素合成的原料少,加之部分碳水化合物用于色素的合成,导致其纤维产量和品质比白色棉低。陆陆杂种纤维中较高的碳水化合物含量可以补偿用于色素合成所消耗的部分碳水化合物含量。可见,天然彩色棉纤维中色素和纤维素的合成在碳水化合物的利用上存在着此消彼长的关系。
3、天然彩色棉纤维细胞氧化还原状态与外源呼吸抑制剂对纤维发育的影响
通过测定白色棉、棕色棉、绿色棉和两个棕色棉杂种纤维细胞发育过程中细胞含水量、pH值、细胞色素c氧化酶活性、NAD~+、NADH、NADP~+、和NADPH的含量及在组培条件下研究了鱼藤酮和硫脲两种呼吸抑制剂对白色棉和绿色棉纤维显色、胚珠鲜重和纤维长度及细胞色素c氧化酶和多酚氧化酶活性的影响。结果表明,绿色棉和棕色棉纤维细胞的显色时间和方式存在明显的差异。棕色棉在30 DPA后以“爆发式”显色;绿色棉则在20 DPA时以“渐进式”逐步使绿色加深,因此,棕色棉和绿色棉纤维中的生理代谢状况是不一样的。白色棉纤维中的水分含量较高,其纤维细胞中的pH值比绿色棉低,但比棕色棉系列棉花高。棕色棉系列纤维细胞中的细胞色素c氧化酶、NAD~+、NADH、NADP~+、和NADPH含量均远远高于白色棉和绿色棉,而且NAD~+或NADP~+含量比NADH或NADPH高,因此,其纤维细胞中的呼吸代谢旺盛;白色棉和绿色棉纤维细胞中的辅酶含量较低,而且两者差异不大。绿色棉在呼吸代谢强度上与白色棉相当,但综合纤维中的色素和纤维素合成两大重要代谢,因此,可以推断绿色棉纤维中大量碳水化合物可能未能被充分利用:相反棕色棉虽然呼吸代谢旺盛,但纤维细胞中的碳水化合物含量少,故造成其纤维素含量比白色棉低,但比绿色棉高;而色素含量比白色棉高,但比绿色棉低。鱼藤酮和硫脲均可抑制白色棉和绿色棉胚珠和纤维细胞的发育,胚珠鲜重减轻,纤维减短,但鱼藤酮的毒害作用更大,而且可使绿色棉纤维显色较对照(绿色棉末用鱼藤酮处理)提早1周;硫脲对胚珠和纤维的发育伤害虽然较小,但可使绿色棉纤维不显色,显然,硫脲对绿色棉纤维的色素代谢具有抑制作用。在鱼藤酮处理下,纤维细胞中的细胞色素c氧化酶的活性迅速降低,而在硫脲各浓度处理下,绿色棉纤维中的多酚氧化酶活性均降低了80%以上。
4、天然彩色棉纤维细胞碳水化合物代谢关键酶活性及基因表达研究
本试验以白色棉、棕色棉、绿色棉和2个棕色棉杂种为材料研究了纤维0-20 DPA发育阶段纤维中酸性和中性转化酶和蔗糖合成酶、蔗糖磷酸合成酶的活性。并且采用RT-PCR和荧光定量RT-PCR的方法对蔗糖合成酶基因的表达量进行了测定。结果表明,棕色棉纤维细胞中的转化酶和蔗糖合成酶最高,绿色棉和白色棉纤维细胞中转化酶和蔗糖合成酶活性均较低相差不大。在2 DPA至20 DPA期间,白色棉和天然彩色棉纤维细胞中的蔗糖合成酶活性高且变化幅度很小,说明纤维细胞发育过程中碳水化合物代谢非常旺盛。白色棉、棕色棉和绿色棉纤维细胞中蔗糖合成酶基因的表达量随纤维发育时期的推进变化不大,但棕色棉纤维细胞中的蔗糖合成酶基因的表达量比白色棉高,绿色棉相对较低。此外,白色棉、棕色棉和绿色棉纤维细胞中的蔗糖合成酶基因的表达量比胚珠和叶片的表达量高。绿色棉纤维细胞中蔗糖合成酶活性和基因表达量低是其碳水化合物利用率低的重要原因。
5、棉花纤维发育特异启动子的克隆及其应用
从棉花基因组DNA中分离到棉花纤维发育特异基因GaRDL 1的启动子,并在其后连接绿色荧光蛋白作为报告基因,采用花粉管通道法转化胚珠。结果表明,在子房注射3天后可在纤维中检测到绿色荧光蛋白,表明该基因启动子在棉花纤维中特异表达。利用反义抑制的方法,在启动子后面连接蔗糖合成酶反义基因,并采用花粉管通道法转化胚珠后,取子房注射3天后的胚珠与纤维的混合物进行蔗糖合成酶活性和碳水化合物含量的测定。结果表明,白色棉、棕色棉和绿色棉纤维中的蔗糖合成酶活性降低,但蔗糖含量因未被蔗糖合成酶分解,因此,蔗糖含量变化不大,同时导致纤维中的葡萄糖和果糖含量降低,从而降低了纤维的碳水化合物利用率。
Low yield and quality of naturally colored cotton fiber is one of the most important problems that is required to be solved in the genetic improvement. During fiber development, besides cellulose synthesis, pigment metabolism is an additional process in naturally colored cotton fiber that differed in white cotton fiber. Pigment synthesis in fiber will consume part of carbohydrate in fiber cells affecting cellulose synthesis for fiber yield and quality. Therefore, in present paper, the molecular and physiological mechanism for the difference of yield and quality between white and naturally colored cotton fibers were analyzed from plant photosynthesis, fiber carbohydrate and pigment content, fiber respiration, and gene function and expression that related with carbohydrate metabolic enzymes aspects according to substrate competition hypothesis. The main results wereshowed as below.
1. Relationship between photosynthetic property and yield and quality of naturally colored cotton fiber
Using brown fiber cotton, green fiber cotton, two brown hybrids, hybrid 1 (the parents were upland cotton) and hybrid 2 (the two parents were upland cotton and sea island cotton), white fiber cotton (control) as plant material, variation of photosynthetic pigment content, chloroplast Hill reaction activity, Mg~(2+)-ATPase and Ca~(2+)-ATPase activity, net photosynthetic rate, leaf starch, sucrose, fructose, and glucose content was determined. Furthermore, correlation and path analysis among yield and quality and these photosynthetic indexes were also analyzed. Result showed that yield ranking of cotton fiber was hybrid 1>white fiber cotton>hybrid 2> brown fiber cotton>green fiber cotton. Yield of green fiber cotton and brown fiber cotton was only 58% and 66% of white fiber cotton, respectively. Although leaf photosynthetic pigment content, net photosynthetic rate and other photosynthetic indexes of green fiber cotton plant was near to white fiber cotton even much higher, its yield was yet the lowest. Lower yield and quality in green fiber cotton was mainly caused by excessively nutritional plant growth, less bolls per plant, and lighter boll mass. However, the reason for low yield and quality of brown fiber cotton was its weak photosynthesis, less physiological activity of leaf chloroplast, and small amount of photosynthetic production. As for the two brown hybrids, photosynthetic ability of hybrid 1 much better than hybrid 2 and yield of hybrid 1 was significantly higher than hybrid 2. However, quality of hybrid 2 was slightly better than hybrid 1. Result of correlation and path analysis among yield and quality and photosynthetic indexes indicated that net photosynthetic rate had the most contribution to yield and the next important was carbohydrate content while had the reverse tendency for fiber length, which was possibly due to its fiber cellular osmotic regulation promoting fiber elongation besides it was used as metabolic substrate. These results indicated that it was feasible for fiber yield and quality improvement through screening for efficiency photosynthesis hybrid combination.
2. Effect of fiber carbohydrate and pigment content on fiber length
Fiber carbohydrate and pigment content, cellulose content, and fiber length dynamic were investigated in white fiber cotton, brown fiber cotton, green fiber cotton, and two brown hybrids during fiber development. Result showed that fiber sucrose content was the highest at 10 DPA and then continuously decreased in white fiber cotton and naturally fiber cotton from the day of anthesis to fiber maturation. Fiber fructose content kept decreasing state from 0 DPA to fiber maturation. However fiber glucose content peaked at 20 DPA, which was consistent with the vigorous stage of the fiber cellulose synthesis both in white and naturally fiber cotton. Fiber carbohydrate content in green fiber cotton and brown hybrid 1 was similar and much higher than other type cotton. However, green fiber cotton had the lowest yield and quality. It was because higher PAL activity leading to large amount of pigment synthesis in green cotton fiber, which would consume much carbohydrate. Thus cellulose synthetic amount reduced. In addition to this reason, low carbohydrate use efficiency was other possibly important reason for low cellulose content because of its low micronaire value, which indicated much carbohydrate was not to be used in cellulose synthesis as fiber maturation. In brown cotton fiber, its lower cellulose content than white cotton fiber was due to low carbohydrate content in fiber and consumption of part carbohydrate for pigment synthesis resulting in substrate reduction. As for hybrid 1, much higher carbohydrate content in fiber could partly make up for the usage of carbohydrate in pigment synthesis. These results demonstrated the substrate ebbing and flowing phenomenon in carbohydrate consumption for pigment and cellulose synthesis.
3. Impact of cellular redox state and exogenous respiratory inhibitors on fiber development
In this section, cellular water content, pH value, cytochrome c oxidase activity, NAD~+, NADH, NADP~+, and NADPH content was monitored as fiber cell development. Furthermore, color display, ovule fresh weight, fiber length, cytochrome c oxidase and polyphenoloxidase activity in green and white cotton fiber were also traced under two respiratory inhibitor treatment, rotenone and thiourea, with in vitro culture condition. Result showed that color display timing and manner was obviously different in green and brown cotton fiber. It displayed as "suddenly" style after 30 DPA in brown cotton fiber while as "increasingly" after 20 DPA in green cotton fiber. Therefore, it could be inferred that their physiological state was different in brown and green cotton fiber. White cotton fiber had much higher water content. However, its pH value was lower than green cotton fiber and higher than brown cotton fiber. Cytochrome c oxidase activity, NAD~+, NADH, NADP~+, and NADPH content in brown serials of cotton fiber was far higher than white and green cotton fiber and NAD~+ or NADP~+ content was much higher than NADH or NADPH demonstrating vigorous respiration in fiber cells. However, coenzyme content in white and green fiber was relatively low and little difference between them. Because green cotton fiber contained two important metabolic systems, which were pigment and cellulose synthesis, it could be inferred lots of carbohydrate might be fully used. Although high activity of respiratory metabolism in brown cotton fiber, cellulose content was higher than green cotton fiber and lower than white cotton fiber while pigment content reversely because of less amount of carbohydrate content in fiber. Ovule and fiber development would be inhibited by both rotenone and thiourea causing ovule fresh weight reduction and fiber shortening. But rotenone was more harmful to ovule and fiber development and color displayed for one week earlier in green cotton fiber. Although thiourea had less effect on ovule and fiber development, it could result in no color display in green cotton fiber indicating pigment metabolism was inhibited by thiourea. Moreover, cytochrome c oxidase activity rapidly decreased in fiber cells after rotenone treatment and polyphenoloxidase activity reduced by more than 80% in green cotton fiber after thiourea treatment.
4. Activity of key enzymes in carbohydrate metabolism in naturally cotton fibers and sucrose synthase gene expression
Carbohydrate metabolic related enzymes, i.e. acidic and neutral invertase, sucrose synthase, and sucrose phosphate synthase, were measured in white, brown, green cotton fiber, and two brown hybrids when fiber was at 0-20 DPA developmental stages. Furthermore, expression amount of sucrose synthase gene was assayed using RT-PCR and real time RT-PCR method. Result showed that activity invertase and sucrose synthase was the highest in brown cotton fiber. These enzymatic activities were similar between green and white cotton fiber. Sucrose synthase activity in white and naturally colored cotton fiber varied little from 2 DPA o 20 DPA revealing high activity of carbohydrate metabolism during cotton fiber development. Furthermore, sucrose synthase gene expression in white, brown, and green cotton fiber altered little during fiber development which was in accordance with enzyme activity variation. However, expression level of sucrose synthase in brown cotton fiber was higher than white cotton fiber. Green cotton fiber had the lowest expression level in fiber cells. As for different organs, fiber cells had much higher expression level than ovule and leaf in white, brown, and green fiber cotton. Low activity of sucrose synthase and gene expression in green cotton fiber was an important reason for its low utilization efficiency of carbohydrate.
5. Molecular cloning of cotton fiber development specific promoter and its application
A cotton fiber development specific promoter of GaRDL 1 gene had been isolated from cotton genomic DNA. The promoter was connected GFP sequentially as report gene and was transformed into ovule by pollen tube pathway. Result showed that GFP could be detected in fiber after 3 days of ovary injection uncovering the specific expression of promoter in fiber. Sucrose synthase gene was reversely ligased after promoter and sucrose synthase activity and carbohydrate content of mixture of ovule and fiber was measured after 3 days of ovary injection using antisense inhibition method. Result showed that sucrose synthase activity reduced in white, brown, and green cotton fiber. However, sucrose content changes little because of its low degradation resulting in low content of glucose and fructose. And thus carbohydrate utilization efficiency was lowered.
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