红壤水分条件对柑橘生理生态要素影响及其作用机理研究
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摘要
中国红壤地区季节性干旱非常严重,土壤水分条件是制约该地区作物和果树生长发育的主要环境因子之一。柑橘在中国栽培面积居世界前列,是中国红壤地区栽培面积最大的水果,是该地区农业的支柱产业和优势产业之一。为探索柑橘在不同红壤水分状况下的生理生态要素变化特征和机理,或为减轻红壤干旱对柑橘生长的伤害和对产量、品质的负面影响,或为红壤地区柑橘果园节水灌溉提供理论依据,本文结合国家“863”高技术研究发展计划农业节水专项(2002AA2Z4331)的实施,于2003-2004年在中国典型红壤地区(江西鹰潭)进行了不同红壤水分条件下柑橘的生理生态反应试验。实验以第四纪红黏土发育的红壤、2年生宫川温州蜜柑(Citrus unshiu Marc.cv.Miyagawa Wase)盆栽幼树(中国科学院红壤生态实验站玻璃温室内)和9龄成年树(中国科学院红壤生态实验站试验示范柑橘园内)为对象,采用土壤水分探头(FDR)实时监测红壤水分含量,控制红壤含水量5个处理:SWC30、SWC45、SWC60、SWC75和SWC90(分别代表红壤相对含水量30%、45%、60%、75%和90%),对柑橘在不同红壤水分条件下的根系、茎和叶生长发育、根系和叶片主要营养元素含量、叶片光合特性、脯氨酸、多胺、活性氧代谢和保护酶活性、果实产量和品质等一系列主要生理生态指标进行实验测定,并对柑橘这些生理生态指标与红壤含水量之间的关系进行了分析。最后,对不同红壤水分条件下柑橘生理生态要素值进行主成分分析,并简要讨论了柑橘生长和生理要素与红壤地区气候环境因子之间的关系。
主要结果如下:
1.红壤在一定的含水量范围内(SWC≤75%),柑橘根系的鲜重、干重、根尖数、总根长、主根长、根平均直径、根体积、根上下扫描面积、根表面积、根不同区域(直径)的总根长和根表面积等根系生长参数以及叶面积、叶周长、叶宽值、树高、新梢长度和横径等根、茎和叶生长指标均随红壤水分增加而增加。其中,依据模拟函数式计算结果,柑橘根体积、总根长、根上下扫描面积、根表面积等生长指标在红壤相对含水量75.5%-79.6%时达最大值。根系和树体高度在红壤相对含水量低于30%、梢长和横径在红壤相对含水量低于45%时受到显著抑制。红壤水分增加(在SWC30%-75%范围内)促进了柑橘根系氮、钾的积累,而降低了磷的含量。其中,根系磷含量与SWC呈显著线性负相关,钾含量与SWC呈显著线性正相关;红壤水分在亏缺或盈余下,柑橘叶片氮含量显著下降;磷、钾含量随红壤水分增加而增加;当SWC≤30%,钙、镁和铁的积累受到了显著抑制。
2.红壤水分影响柑橘叶片叶绿素含量和光合特性。在SWC75处理时,柑橘叶片叶绿素a、叶绿素b、类胡萝卜素含量、净光合速率(Pn)、气孔导度(Gs)、进出气室CO_2的浓度差(△ca)和羧化效率(CE)最高,胞间CO_浓度(Ci)最低。在SWC≤75%时,Pn、△Ca均随红壤水分的增加而显著增加,Pn、△Ca均与SWC呈显著线性正相关,相关系数分别为R~2=0.938 8~*(n=60)和R~2=0.907 2~*(n=60),而Ci变化趋势恰好相反。蒸腾速率(Tr)与SWC(30%-90%)呈极显著线性正相关(R~2:0.9625~(**),n=75)。综合分析Ci、Pn和Gs与红壤水分之间的关系,不同红壤水分条件所引起的柑橘叶片光合速率的变化,在红壤相对含水量低于30%时主要是由非气孔因素所引起的,而高于45%时主要是由气孔因素所引起的。比较光合速率Pn和蒸腾速率Tr与红壤水分含量之间的关系,在红壤相对含水量75%以上时,柑橘叶片蒸腾作用有部分并没有充分用于光合,属无效的奢侈蒸腾。在红壤相对含水量45%-60%时,柑橘水分利用效率(WUE)最高。干旱的SWC30和多水的SWC90处理,WUE则显著降低。
3.红壤水分严重亏缺或盈余影响到柑橘叶片氮代谢次生产物的变化,造成柑橘体内脯氨酸(Pro)和多胺(PAs)等在柑橘体内的积累,使柑橘形成对红壤水分亏缺或盈余等逆境环境的适应性。柑橘叶片游离氨基酸总量和Pro含量随红壤水分的减少而显著增加,游离氨基酸总量(y)与SWC(x)之间呈显著负相关(y=-0.0282x+12.049;R~2=0.852 4~*;n=50),Pro含量(y)在红壤相对含水量低于75%的水分条件下,与SWC(x)呈显著负相关(y=-0.0152x+4.224;R~2=0.860 5~*;n=40)。腐胺(Put)含量在SWC75处理时最低。亚精胺(Spd)含量随红壤水分增加呈抛物线变化,在SWC45处理时含量最高。而精胺(Spm)含量在红壤水分SWC60处理时最高。Spd含量与Put含量之间呈显著正相关。(Spd+Spm)/Put的比值(y)在SWC≤75%的红壤水分条件时,与SWC(x)呈显著线性相关(y=0.0112x+0.173;R~2=0.851 8~*;n=40),柑橘受红壤水分胁迫时产生的PAs对生理起调节作用的主要是Spd与Spm,(Spd+Spm)/Put的比值决定了柑橘受红壤水分胁迫影响的程度。初步提出这个比值可能是柑橘响应红壤水分胁迫程度的一个潜在的敏感度指标。比值越高,柑橘受红壤水分胁迫越轻。这是本实验发现的有关柑橘响应红壤水分变化一个新的生理现象,值得进一步探讨。
4.红壤水分影响了柑橘体内质膜透性以及活性氧含量和保护酶活性的变化。随红壤水分减少,柑橘叶片丙二醛(MDA)含量和质膜透性显著增加,叶片脂质过氧化加重。MDA含量和叶片相对电导率均与SWC呈显著线性负相关,相关系数分别为R~2=0.905 1~*(n=50)和R~2=0.823 6~*(n=50);但柑橘叶片O_2~-生成速率和H_2O_2含量均随红壤水分减少呈下降趋势,而且与柑橘根系和叶片的铁含量显著相关,O_2~-、H_2O_2并不是造成柑橘叶片在红壤干旱胁迫下氧化伤害的主要原因,这是本实验发现柑橘对红壤干旱响应又一个新的生理现象,有待进一步探讨。随着红壤含水量的变化,柑橘叶片抗氧化系统处于动态的平衡中,其中,SOD、POD、APX酶活性在红壤水分亏缺条件下都保持较高的活性,但CAT活性随红壤水分下降却显著降低。
5.红壤水分影响柑橘果实的产量因子和品质指标。红壤水分的增加,柑橘果实生长量显著增加,果实纵/横径比值下降。在SWC≤75%时,柑橘产量,单果重,单果果肉重、果实可食率、果肉含水量等随SWC增加而显著增加,在75%时各指标值最高;果实可溶性固形物、可溶性糖和可滴定酸含量分别与SWC呈显著或极显著线性负相关,相关系数分别为R~2=0.880 7~*(n:50)、R~2=0.717 7~*(n=50)和R~2=0.965 1~(**)(n=50);果实pH值则呈相反趋势,柑橘果实固酸比和糖酸比随SWC的增加而显著增加,果实固酸比与SWC呈显著线性相关(R~2=0.908 6~*,n=50),糖酸比在SWC=75%时达到最大值。红壤水分含量降低可增加柑橘果实可溶性固形物和糖的含量,却降低了果实的适口度。保持红壤含水量60%-75%的范围,即可增加柑橘果实中品质成分的含量,又可提高果实的口感。以柑橘产量、品质和口感而言,柑橘最佳的红壤水分含量为SWC=75%。
6.基于主成分分析,得出红壤水分对柑橘生理生态要素影响相对较大的指标是:树高(y_1)、单果重(y_2)、叶片Fe含量(y_3)、MDA含量(y_4)、Spd含量(y_5)、质膜透性(y_6)、O~(2-)生成速率(y_7)、脯氨酸/游离氨基酸比值(y_8)和柑橘水分利用效率WUE(y_9)。各指标与红壤相对含水量之间(x)的关系如下:
y_1=0.9367x+26.644,R~2=0.928 4~*;
y_2=0.3638x+70.994,R~2=0.898 9~*;
y_3=-2E~(-05)x~2+0.0023x+0.0934,R~2:0.725 0~*;
y_4=-2E~(-05)x+0.004,R~2=0.933 7~(**);
y_5=-0.06x~2+6.7931x-61.917,R~2=0.906 2~(**);
y_6=-0.4152x+14.958,R~2=0.823 6~*;
y_7=-0.0002x~2+0.0278x+0.7546,R~2=0.782 2~*;
y_8=1E~(-05)x~2-0.0017x+0.3754,R~2=0.812 7~*;
y_9=-0.0043x~2+0.5092x-6.3014,R~2=0.910 6~*.
Water content in red soil makes an important role in the production of crop and fruit as one of the most limiting factors in the red soil region of China. Citrus production is the main fruit in the region, and also an important and predominant sector of agriculture whose planting acre stands on the list all over the world.
The objectives of this study were to investigate the effect of soil water content on the characteristics of citrus eco-physiological changes and its mechanism; to lower the negative influence of soil aridity on the growth, output and quality of citrus; further, to provide the theoretical foundation for economic irrigation of citrus production in red soil region of China. So we made the tests of ecophysiology of citrus in the red soil which contain different amount of water in the typical red soil region of China (Yintan, Jiangxi Province) from 2003 to 2004. This project was also put in practice together with the economic irrigation project which was supported by National High Technology Research and Development Programe of China ("863").
Two-year potted plants and 9-year mature satsuma mandarin trees (Citrus unshiu Marc.cv. Miyagawa Wase) were selected as experimental materials, and grown in a red soil developed from the Quaternary red clay. Soil water content was monitored in real time by FDR and controlled to be at 5 different moisture levels(treatments): SWC30, SWC45, SWC60, SWC75 and SWC90 (i.e., soil relative water content as 30%, 45%, 60%, 75% and 90%, respectively). A series of targets of Eco-physiological factors of citrus trees were investigated, such as growth of the root system, stem and leaf; absorption and accumulation of the main nutrient elements; photosynthetic characteristics; content of proline (Pro) and polyamine (PAs); metabolism of reactive oxygen species (ROS) and activities of protect enzymes (APX, CAT, POD and SOD); fruit yield and quality response to soil water contents. The relationship between those eco-physiological factors and soil water content were further analyzed by principal component analysis (PCA). The relationships between citrus growth factors and meteorological-environmental factors were also discussed in this paper.
The main findings of the study are highlighted below:
1. The root growth factors including fresh and dry weight of single citrus root, root tip number, total root length, taproot length, average root diameter, root volume, projection area, surface area, length and surface area in different root regions, and the leaf growth factors including leaf area, leaf perimeter, leaf width, the height of trees, new shoot length, and stem diameter were all increased with the increase of SWC. Among of them, the major root growth factors, such as root volume, total root length, root projection area and root area, were maximized when SWC were from 75.5% to 79.6%. Both root growth and tree height were significantly restrained when SWC was below 30%, and shoot length and stem diameter were also inhibitated when SWC was below 45%. Nitrogen (N) and potassium (K) accumulation in roots was increased while phosphorous (P) content was reduced with the increasing of SWC from 30%-75%. Therefore, it was a negative-linear relation between P content of the roots and SWC while a positive-linear relation between K content and SWC. The N content of citrus leaves was reduced markedly when soil water content was either too low or too high. However, the P and K contents of citrus leaf were increased as the SWC increased. The accumulation of calcium (Ca), magnesium (Mg) and iron (Fe) was negatively affected when SWC was below 30%.
2. Chlorophyll (Chl) content and photosynthetic characteristics were affected by SWC. The content of Chl a, Chl b and carotinoid(Car), net photosynthetic rate(Pn), stomatal conductance(Gs), Difference of CO_2 concentration pass in and out of the air chamber(△Ca) and candarboxylation efficiency(CE) in citrus leaves were maximized while intercellular CO_2 concentrations(Ci) were minimized at 75% SWC. Both Pn and△Ca were increased with the increasing of SWC until 75% (R~2 = 0.938 8*and R~2 = 0.907 2*, respectively). However, Ci decreased as SWC increased. Transpiration rate (Tr) had a significantly positive correlation with SWC (from 30% to 90%, R~2 = 0.962 5**). Analyzing the relationships between Ci, Pn, Gs and SWC, we concluded that Pn variation of citrus leaves resulted from SWC was mainly due to non-stomatal factors when SWC was less 30% and due to stomatal factors if SWC was more than 45%. Part of transpiration was non-effective, in other words, not used for photosynthesis, when SWC was above 75% based on the relationship of Pn and Tr with SWC. The water use efficiency (WUE) of citrus was at the highest when SWC was from 45% to 60%, whereas WUE was markedly reduced under very dry (SWC<30%) and excessive soil water (SWC>90%) conditions.
3. Nitrogen metabolites of citrus leaves were influenced by either little or too much soil water, which resulted in the accumulation of proline (Pro) and polyamine (PAs) in citrus. Total free amino acid (FAA)(y) increased with the decrease of SWC(x)(y=-0.0282x+ 12.049; R~2=0.852 4*, n=50 ). There was a negative relation between Pro(y) and SWC(x) when SWC was below 75% (y=-0.0152x+4.224, R~2=0.860 5*, n=40). Putrescine (Put) content reached the minimum when SWC was 75%. Spermidine (Spd) content changed with the increase of SWC in a parabola shape, and was maximized when SWC was 45%. The spermine (Spm) was maximized at SWC60. There was a significant relationship between Spd(y) and Put(x) (y=0.2404x~2-51.337x +2976.4, R~2=0.858 6*, n=50). When SWC was less than 75%, it had a significantly linear relation between the ratio of (Spd+Spm)/Put(y) and SWC(x) (y=0.0112x+0.173 ,R~2=0.851 8*, n=50). Spd and Spm acted as the main factors in physiological adjustment of PAs, which was produced under the circumstance of soil water stress. And the degree of effect of citrus on soil water stress was determined by the ratio of (Spd+Spm)/Put. The ratio of (Spd+Spm)/Put determined the severity of soil water stress on citrus tree. The higher the ratio, the slighter the soil water stress of the citrus tree was. This physiological phenomenon how citrus trees were affected by soil water regime in the red soil was first discovered by this study, and will be further investigated.
4. Cytoplasm membrane permeability (CMP), ROS and the activities of protective enzymes were affected by soil moisture conditions. As soil water content reducing, malondialdehyde (MDA) and CMP of citrus leaves increased evidently and superoxidation of lipid metabolism was also aggravated. Both MDA content and CMP were significantly correlated with SWC (R~2 = 0.905 l*(n=50) and R~2 = 0.823 6* (n=50) respectively). Superoxide anion radical (?) producing rate and H_2O_2 content all were reduced with the decrease of soil water content, which was related to the Fe content of both citrus roots and leaves. However, this was not the main reason that (?) and H_2O_2 led to the oxidative damage of citrus leaf under the red soil drought conditions. This response of citrus trees to soil drought in the red soil was another new physiological phenomenon found from this experiment and more study is needed to better understand the findings. The anti-oxidation system of citrus leaves was in the homeostasis as SWC was changing. The activities of SOD, POD and APX remained high under the deficit of soil water, while CAT activity decreased rapidly with the decrease of SWC.
5. SWC affected the yield and quality of satsuma mandarin citrus fruit. With the increasing of SWC, fruit yield of the citrus was significantly increased while the ratio of fruit length to diameter decreased. Fruit yields, weight of single fruit, per fruit pulp weight, fruit edible ratio, water content of sarcocarp were all increased with the increase of SWC and reached the maximum at SWC75. Soluble brix, soluble suger and titratable acid of fruit had notable or significantly notable negative-linear relations with SWC (R~2 = 0.880 7*; R~2 = 0.717 7* and R~2=0.965 1**) while pH of fruit was increased as SWC did. Brix-acid ratio of fruit was increased linearly with the increase of SWC (R~2=0.908 6*, n=50) and reached the maximum at SWC75. Suger-acid ratio also increased with SWC. Lowering SWC increased the contents of solube brix and solube suger but reduced the flavor or taste of the fruit of citrus. When SWC was kept in 60% to 75%, quality of satsuma mandarin fruit and the flavor or taste of fruit was all improved. The optimized citrus fruit yields, quality and flavor or taste were achieved at SWC75.
6. Using principal component analysis, it was found that citrus eco-physiological factors influenceded mainly by SWC were tree height (y_1), single fruit weight (y_2), content of Fe (y_3, MDA (y_4), Spd (y_5), CMP (y_6), (?) production rate (y_7), the ratio of Pro to FAA(y_8) and WUE (y_9) of citrus leaves. The relationships between those factors with SWC(x) are listed as following:
主要结果如下:
1.红壤在一定的含水量范围内(SWC≤75%),柑橘根系的鲜重、干重、根尖数、总根长、主根长、根平均直径、根体积、根上下扫描面积、根表面积、根不同区域(直径)的总根长和根表面积等根系生长参数以及叶面积、叶周长、叶宽值、树高、新梢长度和横径等根、茎和叶生长指标均随红壤水分增加而增加。其中,依据模拟函数式计算结果,柑橘根体积、总根长、根上下扫描面积、根表面积等生长指标在红壤相对含水量75.5%-79.6%时达最大值。根系和树体高度在红壤相对含水量低于30%、梢长和横径在红壤相对含水量低于45%时受到显著抑制。红壤水分增加(在SWC30%-75%范围内)促进了柑橘根系氮、钾的积累,而降低了磷的含量。其中,根系磷含量与SWC呈显著线性负相关,钾含量与SWC呈显著线性正相关;红壤水分在亏缺或盈余下,柑橘叶片氮含量显著下降;磷、钾含量随红壤水分增加而增加;当SWC≤30%,钙、镁和铁的积累受到了显著抑制。
2.红壤水分影响柑橘叶片叶绿素含量和光合特性。在SWC75处理时,柑橘叶片叶绿素a、叶绿素b、类胡萝卜素含量、净光合速率(Pn)、气孔导度(Gs)、进出气室CO_2的浓度差(△ca)和羧化效率(CE)最高,胞间CO_浓度(Ci)最低。在SWC≤75%时,Pn、△Ca均随红壤水分的增加而显著增加,Pn、△Ca均与SWC呈显著线性正相关,相关系数分别为R~2=0.938 8~*(n=60)和R~2=0.907 2~*(n=60),而Ci变化趋势恰好相反。蒸腾速率(Tr)与SWC(30%-90%)呈极显著线性正相关(R~2:0.9625~(**),n=75)。综合分析Ci、Pn和Gs与红壤水分之间的关系,不同红壤水分条件所引起的柑橘叶片光合速率的变化,在红壤相对含水量低于30%时主要是由非气孔因素所引起的,而高于45%时主要是由气孔因素所引起的。比较光合速率Pn和蒸腾速率Tr与红壤水分含量之间的关系,在红壤相对含水量75%以上时,柑橘叶片蒸腾作用有部分并没有充分用于光合,属无效的奢侈蒸腾。在红壤相对含水量45%-60%时,柑橘水分利用效率(WUE)最高。干旱的SWC30和多水的SWC90处理,WUE则显著降低。
3.红壤水分严重亏缺或盈余影响到柑橘叶片氮代谢次生产物的变化,造成柑橘体内脯氨酸(Pro)和多胺(PAs)等在柑橘体内的积累,使柑橘形成对红壤水分亏缺或盈余等逆境环境的适应性。柑橘叶片游离氨基酸总量和Pro含量随红壤水分的减少而显著增加,游离氨基酸总量(y)与SWC(x)之间呈显著负相关(y=-0.0282x+12.049;R~2=0.852 4~*;n=50),Pro含量(y)在红壤相对含水量低于75%的水分条件下,与SWC(x)呈显著负相关(y=-0.0152x+4.224;R~2=0.860 5~*;n=40)。腐胺(Put)含量在SWC75处理时最低。亚精胺(Spd)含量随红壤水分增加呈抛物线变化,在SWC45处理时含量最高。而精胺(Spm)含量在红壤水分SWC60处理时最高。Spd含量与Put含量之间呈显著正相关。(Spd+Spm)/Put的比值(y)在SWC≤75%的红壤水分条件时,与SWC(x)呈显著线性相关(y=0.0112x+0.173;R~2=0.851 8~*;n=40),柑橘受红壤水分胁迫时产生的PAs对生理起调节作用的主要是Spd与Spm,(Spd+Spm)/Put的比值决定了柑橘受红壤水分胁迫影响的程度。初步提出这个比值可能是柑橘响应红壤水分胁迫程度的一个潜在的敏感度指标。比值越高,柑橘受红壤水分胁迫越轻。这是本实验发现的有关柑橘响应红壤水分变化一个新的生理现象,值得进一步探讨。
4.红壤水分影响了柑橘体内质膜透性以及活性氧含量和保护酶活性的变化。随红壤水分减少,柑橘叶片丙二醛(MDA)含量和质膜透性显著增加,叶片脂质过氧化加重。MDA含量和叶片相对电导率均与SWC呈显著线性负相关,相关系数分别为R~2=0.905 1~*(n=50)和R~2=0.823 6~*(n=50);但柑橘叶片O_2~-生成速率和H_2O_2含量均随红壤水分减少呈下降趋势,而且与柑橘根系和叶片的铁含量显著相关,O_2~-、H_2O_2并不是造成柑橘叶片在红壤干旱胁迫下氧化伤害的主要原因,这是本实验发现柑橘对红壤干旱响应又一个新的生理现象,有待进一步探讨。随着红壤含水量的变化,柑橘叶片抗氧化系统处于动态的平衡中,其中,SOD、POD、APX酶活性在红壤水分亏缺条件下都保持较高的活性,但CAT活性随红壤水分下降却显著降低。
5.红壤水分影响柑橘果实的产量因子和品质指标。红壤水分的增加,柑橘果实生长量显著增加,果实纵/横径比值下降。在SWC≤75%时,柑橘产量,单果重,单果果肉重、果实可食率、果肉含水量等随SWC增加而显著增加,在75%时各指标值最高;果实可溶性固形物、可溶性糖和可滴定酸含量分别与SWC呈显著或极显著线性负相关,相关系数分别为R~2=0.880 7~*(n:50)、R~2=0.717 7~*(n=50)和R~2=0.965 1~(**)(n=50);果实pH值则呈相反趋势,柑橘果实固酸比和糖酸比随SWC的增加而显著增加,果实固酸比与SWC呈显著线性相关(R~2=0.908 6~*,n=50),糖酸比在SWC=75%时达到最大值。红壤水分含量降低可增加柑橘果实可溶性固形物和糖的含量,却降低了果实的适口度。保持红壤含水量60%-75%的范围,即可增加柑橘果实中品质成分的含量,又可提高果实的口感。以柑橘产量、品质和口感而言,柑橘最佳的红壤水分含量为SWC=75%。
6.基于主成分分析,得出红壤水分对柑橘生理生态要素影响相对较大的指标是:树高(y_1)、单果重(y_2)、叶片Fe含量(y_3)、MDA含量(y_4)、Spd含量(y_5)、质膜透性(y_6)、O~(2-)生成速率(y_7)、脯氨酸/游离氨基酸比值(y_8)和柑橘水分利用效率WUE(y_9)。各指标与红壤相对含水量之间(x)的关系如下:
y_1=0.9367x+26.644,R~2=0.928 4~*;
y_2=0.3638x+70.994,R~2=0.898 9~*;
y_3=-2E~(-05)x~2+0.0023x+0.0934,R~2:0.725 0~*;
y_4=-2E~(-05)x+0.004,R~2=0.933 7~(**);
y_5=-0.06x~2+6.7931x-61.917,R~2=0.906 2~(**);
y_6=-0.4152x+14.958,R~2=0.823 6~*;
y_7=-0.0002x~2+0.0278x+0.7546,R~2=0.782 2~*;
y_8=1E~(-05)x~2-0.0017x+0.3754,R~2=0.812 7~*;
y_9=-0.0043x~2+0.5092x-6.3014,R~2=0.910 6~*.
Water content in red soil makes an important role in the production of crop and fruit as one of the most limiting factors in the red soil region of China. Citrus production is the main fruit in the region, and also an important and predominant sector of agriculture whose planting acre stands on the list all over the world.
The objectives of this study were to investigate the effect of soil water content on the characteristics of citrus eco-physiological changes and its mechanism; to lower the negative influence of soil aridity on the growth, output and quality of citrus; further, to provide the theoretical foundation for economic irrigation of citrus production in red soil region of China. So we made the tests of ecophysiology of citrus in the red soil which contain different amount of water in the typical red soil region of China (Yintan, Jiangxi Province) from 2003 to 2004. This project was also put in practice together with the economic irrigation project which was supported by National High Technology Research and Development Programe of China ("863").
Two-year potted plants and 9-year mature satsuma mandarin trees (Citrus unshiu Marc.cv. Miyagawa Wase) were selected as experimental materials, and grown in a red soil developed from the Quaternary red clay. Soil water content was monitored in real time by FDR and controlled to be at 5 different moisture levels(treatments): SWC30, SWC45, SWC60, SWC75 and SWC90 (i.e., soil relative water content as 30%, 45%, 60%, 75% and 90%, respectively). A series of targets of Eco-physiological factors of citrus trees were investigated, such as growth of the root system, stem and leaf; absorption and accumulation of the main nutrient elements; photosynthetic characteristics; content of proline (Pro) and polyamine (PAs); metabolism of reactive oxygen species (ROS) and activities of protect enzymes (APX, CAT, POD and SOD); fruit yield and quality response to soil water contents. The relationship between those eco-physiological factors and soil water content were further analyzed by principal component analysis (PCA). The relationships between citrus growth factors and meteorological-environmental factors were also discussed in this paper.
The main findings of the study are highlighted below:
1. The root growth factors including fresh and dry weight of single citrus root, root tip number, total root length, taproot length, average root diameter, root volume, projection area, surface area, length and surface area in different root regions, and the leaf growth factors including leaf area, leaf perimeter, leaf width, the height of trees, new shoot length, and stem diameter were all increased with the increase of SWC. Among of them, the major root growth factors, such as root volume, total root length, root projection area and root area, were maximized when SWC were from 75.5% to 79.6%. Both root growth and tree height were significantly restrained when SWC was below 30%, and shoot length and stem diameter were also inhibitated when SWC was below 45%. Nitrogen (N) and potassium (K) accumulation in roots was increased while phosphorous (P) content was reduced with the increasing of SWC from 30%-75%. Therefore, it was a negative-linear relation between P content of the roots and SWC while a positive-linear relation between K content and SWC. The N content of citrus leaves was reduced markedly when soil water content was either too low or too high. However, the P and K contents of citrus leaf were increased as the SWC increased. The accumulation of calcium (Ca), magnesium (Mg) and iron (Fe) was negatively affected when SWC was below 30%.
2. Chlorophyll (Chl) content and photosynthetic characteristics were affected by SWC. The content of Chl a, Chl b and carotinoid(Car), net photosynthetic rate(Pn), stomatal conductance(Gs), Difference of CO_2 concentration pass in and out of the air chamber(△Ca) and candarboxylation efficiency(CE) in citrus leaves were maximized while intercellular CO_2 concentrations(Ci) were minimized at 75% SWC. Both Pn and△Ca were increased with the increasing of SWC until 75% (R~2 = 0.938 8*and R~2 = 0.907 2*, respectively). However, Ci decreased as SWC increased. Transpiration rate (Tr) had a significantly positive correlation with SWC (from 30% to 90%, R~2 = 0.962 5**). Analyzing the relationships between Ci, Pn, Gs and SWC, we concluded that Pn variation of citrus leaves resulted from SWC was mainly due to non-stomatal factors when SWC was less 30% and due to stomatal factors if SWC was more than 45%. Part of transpiration was non-effective, in other words, not used for photosynthesis, when SWC was above 75% based on the relationship of Pn and Tr with SWC. The water use efficiency (WUE) of citrus was at the highest when SWC was from 45% to 60%, whereas WUE was markedly reduced under very dry (SWC<30%) and excessive soil water (SWC>90%) conditions.
3. Nitrogen metabolites of citrus leaves were influenced by either little or too much soil water, which resulted in the accumulation of proline (Pro) and polyamine (PAs) in citrus. Total free amino acid (FAA)(y) increased with the decrease of SWC(x)(y=-0.0282x+ 12.049; R~2=0.852 4*, n=50 ). There was a negative relation between Pro(y) and SWC(x) when SWC was below 75% (y=-0.0152x+4.224, R~2=0.860 5*, n=40). Putrescine (Put) content reached the minimum when SWC was 75%. Spermidine (Spd) content changed with the increase of SWC in a parabola shape, and was maximized when SWC was 45%. The spermine (Spm) was maximized at SWC60. There was a significant relationship between Spd(y) and Put(x) (y=0.2404x~2-51.337x +2976.4, R~2=0.858 6*, n=50). When SWC was less than 75%, it had a significantly linear relation between the ratio of (Spd+Spm)/Put(y) and SWC(x) (y=0.0112x+0.173 ,R~2=0.851 8*, n=50). Spd and Spm acted as the main factors in physiological adjustment of PAs, which was produced under the circumstance of soil water stress. And the degree of effect of citrus on soil water stress was determined by the ratio of (Spd+Spm)/Put. The ratio of (Spd+Spm)/Put determined the severity of soil water stress on citrus tree. The higher the ratio, the slighter the soil water stress of the citrus tree was. This physiological phenomenon how citrus trees were affected by soil water regime in the red soil was first discovered by this study, and will be further investigated.
4. Cytoplasm membrane permeability (CMP), ROS and the activities of protective enzymes were affected by soil moisture conditions. As soil water content reducing, malondialdehyde (MDA) and CMP of citrus leaves increased evidently and superoxidation of lipid metabolism was also aggravated. Both MDA content and CMP were significantly correlated with SWC (R~2 = 0.905 l*(n=50) and R~2 = 0.823 6* (n=50) respectively). Superoxide anion radical (?) producing rate and H_2O_2 content all were reduced with the decrease of soil water content, which was related to the Fe content of both citrus roots and leaves. However, this was not the main reason that (?) and H_2O_2 led to the oxidative damage of citrus leaf under the red soil drought conditions. This response of citrus trees to soil drought in the red soil was another new physiological phenomenon found from this experiment and more study is needed to better understand the findings. The anti-oxidation system of citrus leaves was in the homeostasis as SWC was changing. The activities of SOD, POD and APX remained high under the deficit of soil water, while CAT activity decreased rapidly with the decrease of SWC.
5. SWC affected the yield and quality of satsuma mandarin citrus fruit. With the increasing of SWC, fruit yield of the citrus was significantly increased while the ratio of fruit length to diameter decreased. Fruit yields, weight of single fruit, per fruit pulp weight, fruit edible ratio, water content of sarcocarp were all increased with the increase of SWC and reached the maximum at SWC75. Soluble brix, soluble suger and titratable acid of fruit had notable or significantly notable negative-linear relations with SWC (R~2 = 0.880 7*; R~2 = 0.717 7* and R~2=0.965 1**) while pH of fruit was increased as SWC did. Brix-acid ratio of fruit was increased linearly with the increase of SWC (R~2=0.908 6*, n=50) and reached the maximum at SWC75. Suger-acid ratio also increased with SWC. Lowering SWC increased the contents of solube brix and solube suger but reduced the flavor or taste of the fruit of citrus. When SWC was kept in 60% to 75%, quality of satsuma mandarin fruit and the flavor or taste of fruit was all improved. The optimized citrus fruit yields, quality and flavor or taste were achieved at SWC75.
6. Using principal component analysis, it was found that citrus eco-physiological factors influenceded mainly by SWC were tree height (y_1), single fruit weight (y_2), content of Fe (y_3, MDA (y_4), Spd (y_5), CMP (y_6), (?) production rate (y_7), the ratio of Pro to FAA(y_8) and WUE (y_9) of citrus leaves. The relationships between those factors with SWC(x) are listed as following:
引文
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