黑龙江省桑树氮肥运筹及光合调控机理的研究
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摘要
黑龙江省地处我国的高寒地区,主要栽培在工矿废弃荒地、山地、田间地头以及贫瘠土壤。国家“东桑西移,南蚕北移”工程及有关优惠政策的实施,加快了北方寒冷地区蚕桑业的快速发展。桑树栽培由田问地头转移到大田栽培。黑龙江省是我国最适宜发展蚕桑生产的地区之一,桑蚕业已成为黑龙江省西部干旱、半干旱地区脱贫致富的支柱产业,尤其是桑树品种“龙桑一号”的培育出世更加速了该地区桑蚕产业的迅速发展。但是,在实际生产中,为了获得较高的桑树叶片产量,一些蚕农未能根据土壤养分状况和桑树需肥规律而无限制地加大氮肥的施用量,这不但不能提高桑树的产叶量,反而加大了投入成本,污染环境,致使养蚕质量下降。桑树是以叶片为收获对象的作物,其栽培的主要目的是获得较高的叶片产量,而氮肥是非常重要的因素。为此,本论文从黑龙江省桑树栽培过程中的氮肥运筹出发,通过多年(2009—2012)多点(哈尔滨、阿城、佳木斯)的试验研究,在确定主栽桑树品种的氮素用量范围的基础上,探讨桑树对氮素形态(硝态氮和铵态氮)的利用,通过大田试验和水培试验相结合的方法,从物质生产的基础一光合作用的光反应和暗反应之间的协同作用角度,深入分析桑树对氮素的响应特性。结果如下：
1.2009-2012年在大田条件下研究了不同供氮(纯氮)水平(0kg·hm-2、48.3kg·hm-2、96.6、144.9kg·hm-2、193.2kg·hm-2和241.5kg·hm-2)和(0kg·hm-2,13.8kg·hm-2,27.6kg·hm-2,41.4kg·hm-2,55.2kg·hm-2,69kg·hm-2)分别对“龙桑一号”和“青龙”桑叶片产量及其生理特性的影响。结果表明：当“龙桑一号”施氮量为0～193.2kg·hm-2时和“青龙”桑施氮量为0～55.2kg·hm-2,桑树单株枝条数、单枝叶片数、单叶面积、单叶质量和单株产叶量随着供氮水平的增加而增加,而当施氮量分别达到241.5kg·hm-2和69kg·hm-2时,桑树单叶面积、叶片质量和单株产叶量反而降低；施氮量在0～144.9kg·hm-2和0～41.4kg·hm-2范围内,桑树叶片中的全氮(N)和全磷(P)含量随着施氮量的增加而提高,而当施氮量超过144.9kg·hm-2和41.4kg·hm-2时,两利桑树叶片中的全氮和全磷含量随着施氮量的增加却降低,这说明施氮量过高反而降低了桑树对N和P的吸收；随着供氮水平的增加,桑树叶片中的叶绿素含量、可溶性蛋白和可溶性糖含量随着施氮量的增加而增加。从氮肥对桑树产叶量及其生理特性的影响情况来看,桑树“龙桑一号”和“青龙”桑田间的最佳施氮量分别为144.9～193.2k·-hm-2和41.4～55.2kg·hm-2。四年大田试验中氮肥与桑树产叶量(Y)的偏最小二乘回归模型如下：
y=1552.3484111+2.702975x1+58.083982x2+9.429218x3-0.000561x1*x1-0.672614x2*x2-0.017726x3*x3-0.036371x1*x2-0.005904x1*x3-0.109191x2*x3
y=1450.0240821+2.983592x1+54.401789x2+9.645707x3-0.001892x1*x1-0.579893x2*x2-0.018230x3*x3-0.032356x1*x2-0.005737x1*x3-0.102818x2*x3
y=1490.3531551+2.752968x1+51.050728x2+8.862974x3-0.001686x1*x1-0.544203x2*x2-0.016403x3*x3-0.030354x1*x2-0.005270x1*x3-0.094480x2*x3
y=608.0876801+4.590733x1+37.395394x2+5.491631x3-0.003846x1*x1-0.483353x2*x2-0.010424x3*x3-0.070032x1*x2-0.010284x1*x3-0.070982x2*x3
2.通过水培方式研究了等氮条件下铵态氮和硝态氮两种形态氮源及其配比对青龙桑幼苗生长和光合特性的影响。结果表明,桑树幼苗在单一硝态氮或单一铵态氮条件下,植株高度、叶片数、叶片面积和根系长度均低于铵态氮和硝态氮配合施用下,桑树叶片和根系生物量的变化也呈现类似趋势。铵态氮和硝态氮摩尔浓度比例为50：50和25：75时桑树幼苗生长和生物量最高,而当铵态氮和硝态氮比例为25：75时桑树净光合速率(Pn)、气孔导度(Gs)和水分利用效率(WUE)高于其它处理,单一硝态氮或单一铵态氮处理降低了桑树表观量子效率(AQE),提高了桑树叶片的光补偿点(LCP)。以上说明桑树是一种偏硝性的植物,以铵态氮和硝态氮比例为50：50～25：75之间最适合。
3.不同氮素形态营养相比较,NH4+和NO3-配比处理的桑树叶片的叶绿素含量(Chl)、光合速率(Pn),NO3-处理次之,NH4+处理较低。各氮素处理的气孔导度与对照相比差异较小。氮素营养使叶片胞间CO2浓度(Ci)降低。与NH4+相比,N03-提高了叶片的光呼吸速率、光呼吸速率与光合速率比值。NH4+和NO3-配比处理叶片光合速率的提高与其较高的Rubisco活力、光合电子传递活性的改善有关。
4.硝态氮和铵态氮作为氮素的两种形态,有着不同的生理生化代谢途径,植物在以硝态氮为氮源的情况下生长良好,而在铵态氮为氮源的情况下则出现“铵盐毒害”现象,被称为“喜硝植物”；也有植物对铵态氮有较高的适应性,即使以铵态氮作为唯一氮源也基本不影响其生长状态,被称为“喜铵植物”。为了说明桑树是那一种类型,从两种氮素形态对光合电子流分配的影响角度来探明不同氮素形态对桑树生理生化产生不同影响的机理及硝酸盐代谢在耗散过剩光能中的光保护作用机制。
1)以铵态氮或高比例的铵态氮作为氮源时,桑树生长受到明显抑制,表现在：地上部和根部生物量显著下降,其中根部受到的抑制更加明显。与硝态氮相比,铵态氮导致桑树净光合速率(Pn)显著降低,其直接原因可能是非气孔因素。
2)铵态氮培养下桑树PSⅡ总电子流JF(PSⅡ、用于碳还原的电子流J0(PCR)和用于光呼吸的电子流Jc(PCO)均显著低于硝态氮培养的植株。铵态氮培养下的桑树ΦPSⅡ显著降低,且伴随着qP的显著下降,Fv'/Fm'基本不变,说明铵态氮引起的光合电子流降低的原因是开放光系统Ⅱ(PSⅡ)反应中心数量的减少。此外,铵态氮导致桑树PSⅡ总电子流下降,客观上需要其他形式光能耗散途径的增强,以保护光系统免于光抑制的破坏。
5.通过不同抗逆能力的桑树品种“龙桑一号”(抗逆能力弱)与“青龙”桑(抗逆能力强)叶片的光合电子传递及激发能利用分配对氮素响应的结果表明,施氮可提高不同品种桑树叶片天线色素吸收光能的能力,虽然氮素不能改变激发能在光合碳还原(PCR)和光合碳氧化(PCO)之间的分配比例,但可提高PSⅡ总电子传递速率(JF)和Pn。氮素对叶片PSⅡ反应中心活性有影响,而不同抗性品种之间亦有差别,说明施氮可改善桑树叶片热耗散和光化学反应对激发能的竞争关系,从而增强光合机构的自我保护能力。
综上所述,从氮素和氮素形态调控桑树生长机理方面出发,以桑树叶片光合电子传递和吸收光能分配特性为线索,揭示了氮素,尤其是氮素形态在桑树体内代谢过程中需要的还原力(NADPH或NADH)的来源,阐明了硝态氮还原为铵态氮需要的还原力大部分来自于光合电子传递链(PSⅠ侧),但是,铵态氮在植物根部同化为氨基酸,其还原力主要来自呼吸作用,从而探明在光能过剩的情况下光合链电子受体库有效消耗了部分过剩激发能,减轻了光抑制对光系统的破坏,并解释了长期以来人们一直争议的硝酸盐还原对光破坏防御机制的贡献,有助于我们进一步理解植物的光破坏防御机制。
Heilongjiang Province is located in the alpine region of our country, the main cultivated in the mining waste wasteland, mountains, the fields, and poor soil. Much more national preferential policies had been to speed up the development of sericulture industry in northern cold regions. Mulberry cultivation farms to transfer to field cultivation. Heilongjiang Province is one of the most suitable for the development of sericulture production areas in China, silkworm has become the pillar industries in the western arid, semi-arid areas of Heilongjiang Province, especially the cultivation of mulberry varieties "Long Sang NO.1" born accelerate the region the rapid development of the silkworm industry. However, in the actual production, in order to obtain a higher yield of mulberry leaves, some silkworm breeder fails to soil nutrient status and mulberry Fertilizer Demand Rule of unlimited increase the amount of nitrogen fertilizer, which will not only fail to improve the mulberry leaf yield, but increased input costs, environmental pollution, resulting in the decline in the quality of sericulture. Leaves of mulberry are harvested object crop cultivation its main purpose is to obtain high yield blade, while nitrogen is a very important factor. To this end, the departure from Nitrogen Fertilizer in Heilongjiang Province mulberry cultivation process, through the years (2009-2012) multi-point (Harbin, Cheng, Jiamusi) test in determining the nitrogen amount in the range of the main cultivated varieties of mulberry based on the use of mulberry nitrogen forms (nitrate and ammonium nitrogen) through field trials and hydroponics method of combining production from the material-the light reactions of photosynthesis and dark reaction between synergy angle, in-depth analysis of the response characteristics of the mulberry nitrogen. The results are as follows:
1. In order to research the best appropriate N application levels of "Longsang No.1" and "qinglong", and support a optimum guide for mulberry peasant, this study was carried out to investigate the effects of different N application levels (0,48.3,96.6,144.9,193.2and241.5kg·hm-2pure nitrogen) and (0kg·hm-2,13.8kg·hm-2,27.6kg·hm-2,41.4kg·hm-2,55.2kg·hm-2,69kg·hm-2) on growth and physiological characteristics of mulberry grown in the field using mulberry variety Longsang No.l and "qinglong" as experimental material. The results showed that leaf number, branch number, leaf area, leaf weight and leaf yield per plant increased with nitrogen application levels in0～193.2and kg·hm-20～55.2kg·hm-2, while the nitrogen application levels in241.5kg·hm-2and69kg·hm-2, the leaf area, leaf weight and leaf yield per plant reduced. Total nitrogen and total phosphorus contents in leaves of mulberry increased with nitrogen application levels in0～144.9kg·hm-2and41.4kg·hm-2, but over nitrogen application levels of that the trend is opposite, the indicating that higher nitrogen can be reduce the absorption of N and P. Chlorophyll, soluble protein contents and soluble sugar contents in leaves of mulberry increased with nitrogen application levels increased. In sum, by the N application levels on the growth and physiological characteristics of mulberry, the best N application levels in the field of "Long sang NO.1" and "qinglong" is144.9～241.5kg·hm-2and41.4～55.2kg·hm-2respectively. Nitrogen fertilizer and mulberry leaf yield amount (Y) in a four-year field experiment partial least squares regression model are as follows:
y=1552.3484111+2.702975x1+58.083982x2+9.429218x3-0.000561x1*x1-0.672614x2*x2-0.017726x3*x3-0.036371x*x2-0.005904x1*x3-0.109191x2*x3
y=1450.0240821+2.983592x1+54.401789x2+9.645707x3-0.001892x1*x1-0.579893x2*x2-0.018230x3*x3-0.032356x1*x2-0.005737x1*x3-0.102818x2*x3
y=1490.3531551+2.752968x1+51.050728x2+8.862974x3-0.001686x1*x1-0.544203x2*x2-0.016403x3*x3-0.030354x1*x2-0.005270x1*x3-0.094480x2*x3
y=608.0876801+4.590733x1+37.395394x2+5.491631x3-0.003846x1*x1-0.483353x2*x2-0.010424x3*x3-0.070032x1*x2-0.010284x1*x3-0.070982x2*x3
2. The effects of different proportions of nitrate and ammonium nitrogen applied to mulberry (Morus alba L.) on plant growth and photosynthetic characteristics of mulberry seedlings cultured in hydroponics were investigated under the same nitrogen amount using forage mulberry variety "Qinglong" as experimental material. Plant height, leaf number, leaf area and root length of mulberry seedlings grown in hydroponics of single ammonium or nitrate nitrogen source were lower than that of proportion solution of ammonium and nitrate nitrogen, leaf and root biomass of mulberry seedlings grown in hydroponics was similar trends. Plant growth and biomass in mulberry seedlings were highest while the proportions of nitrate and ammonium nitrogen were50:50and25:75. Net photosynthetic rate (Pn), stomatal conductance (Gs) and water use efficiency (WUE) in leaves of mulberry seedlings cultured in25:75proportion solution of ammonium and nitrate nitrogen were higher than that of other proportion treatments. Single ammonium or nitrate nitrogen source improved the apparent quantum efficiency (AQE) and decreased light compensation point (LCP) in leaves of mulberry seedlings. It concluded that forage mulberry varieties "Qinglong" prefer nitrate nitrogen, and the proportions of nitrate and ammonium nitrogen adapted mulberry growth were between50:50and25:75.
3. In three different nitrogen treatments, the chlorophyll content (Chl), photosynthetic
Rate (Pn), RuBPcase initial activity, RuBPcase content, mesophyll conductance (gm), carbonic Anhydrase (CA) activity, PS1activity and PS1activity of leaves were the highest in the mixture treatment of NH4++NO3-, then were in the NO3-treatment and the lowest were in NH4+treatment. No evident differences were found in RuBPcase specific activity among treatments. There was little difference on stomatol conductance (Gs) between different treatments and CK. Nitrogen nutritions decreased the intercellular CO2concentration (Ci). There were similarly Changing trends between gm, CA activity and photosynthetic rate (Pn) in different treatments, Which showed the conductivity of CO2in liquid part of mesophyll cell having probably important Influence on photosynthesis of leaves. Compared to NH4+treatment, NO3-treatment increased the photorespiration rate, the ratio of photorespiration rate to photosynthetic rate and nitrate reduetase (NR) activity in wheat leaves. The increase of photosynthetic rate in mixture treatment NH4++NO3-was mainly correlated to the higher Rubisco activity, higher photosynthetic electron conductivity and the improvement of CO2conductivity in liquid phase of mesophall cell.
4. Nitrogen plays an important role throughout the whole life of all living organisms. Nitrate and ammonium are the major forms of nitrogen available to plants. Though both forms can be utilized, great differences in plant growth are found among various plants. Since1950s many research efforts have been directed toward unraveling the causes and mechanisms of NH4+toxicity as well as the relationship of nitrate and photoprotection in plants, while present knowledge is far from complete. It is kown that most of the energy for nitrate assimilation in a cell derives from photosynthesis. Recently, nitrate reduction has been suggested to be involved in the photo protection mechanism. However, our present understanding of this role of nitrate is poor. This research focuses on the difference in allocation. The results are as follows:
1) The effects of different nitrogen forms on biomass and gas exchange of mulberry and rice were studied. Ammonium significantly decreased both shoot biomass and root biomass of mulberry, especially root biomass. Photosynthetie rate (Pn) of mulberry under ammonium condition was also significantly lower than those under nitrate condition while stomatal conductance and intracellular CO2concentration were higher. Thus, decreased Pn could be attributed to stomatal limitation. In comparison, there were no significant differences in biomass and gas exchange between rice cultured under nitrate or ammonium condition.
2) The effects of different nitrogen forms on photosynthetic electron allocation and nitrate reduetase activity were investigated in mulberry and rice plants. When cultured with ammonium as nitrogen source, mulberry plants had lower JF(PS Ⅱ)), Jo(PCR), JC(PCO. The effects of different nitrogen forms on chlorophyll fluorescence and xanthophyll cycle activity were studied in mulberry and rice plants. There was no significant difference in either chlorophyll fluorescence or xanthophyll cycle activity between nitrate and ammonium feed rice plants.
5. The responses of photosynthetic electron transport and excitation energy distribution in winter wheat leaves at the jointing stage to nitrogen levels were studied in the field experiment, both leaf gas exchange parameter and chlorophyll fluorescence were measured. The light energy absorbed by antenna pigment of both cultivars increased with the treatment of nitrogen. Although nitrogen could not obviously change the distribution proportion of photosynthetic carbon reduction and photosynthetic carbon oxidation, nitrogen could increase JF of PSII and photosynthetic rate. Nitrogen could affect PSII activities of both cultivars, and there were significant differences between the two cultivars. Nitrogen application improved the competition between heat dissipation and photo-chemic reaction and enhanced the self-protection ability of photosynthetic apparatus.
In summary, the regulation mechanism of growth of mulberry trees from nitrogen and nitrogen forms, the mulberry leaf photosynthetic electron transport and absorption of light energy distribution characteristics for clues to reveal the nitrogen, especially of nitrogen forms in Mulberry metabolism processsource of reducing power (NADPH or NADH) to clarify the reducing power of nitrate reduction to ammonium nitrogen needs from the photosynthetic electron transport chain (PS I side), but the ammonium nitrogen assimilation into amino acids in plant roots, reducing power mainly from respiration, which proved effective consumption of energy excess the photosynthetic chain electron acceptor library part of the excess excitation energy, thereby reducing the destruction of photoinhibition of photosystem and explain long beencontroversial reduction of nitrate to the contribution of the optical destruction defense mechanism, to help us further understand plant Photoprotective Mechanisms.
1.2009-2012年在大田条件下研究了不同供氮(纯氮)水平(0kg·hm-2、48.3kg·hm-2、96.6、144.9kg·hm-2、193.2kg·hm-2和241.5kg·hm-2)和(0kg·hm-2,13.8kg·hm-2,27.6kg·hm-2,41.4kg·hm-2,55.2kg·hm-2,69kg·hm-2)分别对“龙桑一号”和“青龙”桑叶片产量及其生理特性的影响。结果表明：当“龙桑一号”施氮量为0～193.2kg·hm-2时和“青龙”桑施氮量为0～55.2kg·hm-2,桑树单株枝条数、单枝叶片数、单叶面积、单叶质量和单株产叶量随着供氮水平的增加而增加,而当施氮量分别达到241.5kg·hm-2和69kg·hm-2时,桑树单叶面积、叶片质量和单株产叶量反而降低；施氮量在0～144.9kg·hm-2和0～41.4kg·hm-2范围内,桑树叶片中的全氮(N)和全磷(P)含量随着施氮量的增加而提高,而当施氮量超过144.9kg·hm-2和41.4kg·hm-2时,两利桑树叶片中的全氮和全磷含量随着施氮量的增加却降低,这说明施氮量过高反而降低了桑树对N和P的吸收；随着供氮水平的增加,桑树叶片中的叶绿素含量、可溶性蛋白和可溶性糖含量随着施氮量的增加而增加。从氮肥对桑树产叶量及其生理特性的影响情况来看,桑树“龙桑一号”和“青龙”桑田间的最佳施氮量分别为144.9～193.2k·-hm-2和41.4～55.2kg·hm-2。四年大田试验中氮肥与桑树产叶量(Y)的偏最小二乘回归模型如下：
y=1552.3484111+2.702975x1+58.083982x2+9.429218x3-0.000561x1*x1-0.672614x2*x2-0.017726x3*x3-0.036371x1*x2-0.005904x1*x3-0.109191x2*x3
y=1450.0240821+2.983592x1+54.401789x2+9.645707x3-0.001892x1*x1-0.579893x2*x2-0.018230x3*x3-0.032356x1*x2-0.005737x1*x3-0.102818x2*x3
y=1490.3531551+2.752968x1+51.050728x2+8.862974x3-0.001686x1*x1-0.544203x2*x2-0.016403x3*x3-0.030354x1*x2-0.005270x1*x3-0.094480x2*x3
y=608.0876801+4.590733x1+37.395394x2+5.491631x3-0.003846x1*x1-0.483353x2*x2-0.010424x3*x3-0.070032x1*x2-0.010284x1*x3-0.070982x2*x3
2.通过水培方式研究了等氮条件下铵态氮和硝态氮两种形态氮源及其配比对青龙桑幼苗生长和光合特性的影响。结果表明,桑树幼苗在单一硝态氮或单一铵态氮条件下,植株高度、叶片数、叶片面积和根系长度均低于铵态氮和硝态氮配合施用下,桑树叶片和根系生物量的变化也呈现类似趋势。铵态氮和硝态氮摩尔浓度比例为50：50和25：75时桑树幼苗生长和生物量最高,而当铵态氮和硝态氮比例为25：75时桑树净光合速率(Pn)、气孔导度(Gs)和水分利用效率(WUE)高于其它处理,单一硝态氮或单一铵态氮处理降低了桑树表观量子效率(AQE),提高了桑树叶片的光补偿点(LCP)。以上说明桑树是一种偏硝性的植物,以铵态氮和硝态氮比例为50：50～25：75之间最适合。
3.不同氮素形态营养相比较,NH4+和NO3-配比处理的桑树叶片的叶绿素含量(Chl)、光合速率(Pn),NO3-处理次之,NH4+处理较低。各氮素处理的气孔导度与对照相比差异较小。氮素营养使叶片胞间CO2浓度(Ci)降低。与NH4+相比,N03-提高了叶片的光呼吸速率、光呼吸速率与光合速率比值。NH4+和NO3-配比处理叶片光合速率的提高与其较高的Rubisco活力、光合电子传递活性的改善有关。
4.硝态氮和铵态氮作为氮素的两种形态,有着不同的生理生化代谢途径,植物在以硝态氮为氮源的情况下生长良好,而在铵态氮为氮源的情况下则出现“铵盐毒害”现象,被称为“喜硝植物”；也有植物对铵态氮有较高的适应性,即使以铵态氮作为唯一氮源也基本不影响其生长状态,被称为“喜铵植物”。为了说明桑树是那一种类型,从两种氮素形态对光合电子流分配的影响角度来探明不同氮素形态对桑树生理生化产生不同影响的机理及硝酸盐代谢在耗散过剩光能中的光保护作用机制。
1)以铵态氮或高比例的铵态氮作为氮源时,桑树生长受到明显抑制,表现在：地上部和根部生物量显著下降,其中根部受到的抑制更加明显。与硝态氮相比,铵态氮导致桑树净光合速率(Pn)显著降低,其直接原因可能是非气孔因素。
2)铵态氮培养下桑树PSⅡ总电子流JF(PSⅡ、用于碳还原的电子流J0(PCR)和用于光呼吸的电子流Jc(PCO)均显著低于硝态氮培养的植株。铵态氮培养下的桑树ΦPSⅡ显著降低,且伴随着qP的显著下降,Fv'/Fm'基本不变,说明铵态氮引起的光合电子流降低的原因是开放光系统Ⅱ(PSⅡ)反应中心数量的减少。此外,铵态氮导致桑树PSⅡ总电子流下降,客观上需要其他形式光能耗散途径的增强,以保护光系统免于光抑制的破坏。
5.通过不同抗逆能力的桑树品种“龙桑一号”(抗逆能力弱)与“青龙”桑(抗逆能力强)叶片的光合电子传递及激发能利用分配对氮素响应的结果表明,施氮可提高不同品种桑树叶片天线色素吸收光能的能力,虽然氮素不能改变激发能在光合碳还原(PCR)和光合碳氧化(PCO)之间的分配比例,但可提高PSⅡ总电子传递速率(JF)和Pn。氮素对叶片PSⅡ反应中心活性有影响,而不同抗性品种之间亦有差别,说明施氮可改善桑树叶片热耗散和光化学反应对激发能的竞争关系,从而增强光合机构的自我保护能力。
综上所述,从氮素和氮素形态调控桑树生长机理方面出发,以桑树叶片光合电子传递和吸收光能分配特性为线索,揭示了氮素,尤其是氮素形态在桑树体内代谢过程中需要的还原力(NADPH或NADH)的来源,阐明了硝态氮还原为铵态氮需要的还原力大部分来自于光合电子传递链(PSⅠ侧),但是,铵态氮在植物根部同化为氨基酸,其还原力主要来自呼吸作用,从而探明在光能过剩的情况下光合链电子受体库有效消耗了部分过剩激发能,减轻了光抑制对光系统的破坏,并解释了长期以来人们一直争议的硝酸盐还原对光破坏防御机制的贡献,有助于我们进一步理解植物的光破坏防御机制。
Heilongjiang Province is located in the alpine region of our country, the main cultivated in the mining waste wasteland, mountains, the fields, and poor soil. Much more national preferential policies had been to speed up the development of sericulture industry in northern cold regions. Mulberry cultivation farms to transfer to field cultivation. Heilongjiang Province is one of the most suitable for the development of sericulture production areas in China, silkworm has become the pillar industries in the western arid, semi-arid areas of Heilongjiang Province, especially the cultivation of mulberry varieties "Long Sang NO.1" born accelerate the region the rapid development of the silkworm industry. However, in the actual production, in order to obtain a higher yield of mulberry leaves, some silkworm breeder fails to soil nutrient status and mulberry Fertilizer Demand Rule of unlimited increase the amount of nitrogen fertilizer, which will not only fail to improve the mulberry leaf yield, but increased input costs, environmental pollution, resulting in the decline in the quality of sericulture. Leaves of mulberry are harvested object crop cultivation its main purpose is to obtain high yield blade, while nitrogen is a very important factor. To this end, the departure from Nitrogen Fertilizer in Heilongjiang Province mulberry cultivation process, through the years (2009-2012) multi-point (Harbin, Cheng, Jiamusi) test in determining the nitrogen amount in the range of the main cultivated varieties of mulberry based on the use of mulberry nitrogen forms (nitrate and ammonium nitrogen) through field trials and hydroponics method of combining production from the material-the light reactions of photosynthesis and dark reaction between synergy angle, in-depth analysis of the response characteristics of the mulberry nitrogen. The results are as follows:
1. In order to research the best appropriate N application levels of "Longsang No.1" and "qinglong", and support a optimum guide for mulberry peasant, this study was carried out to investigate the effects of different N application levels (0,48.3,96.6,144.9,193.2and241.5kg·hm-2pure nitrogen) and (0kg·hm-2,13.8kg·hm-2,27.6kg·hm-2,41.4kg·hm-2,55.2kg·hm-2,69kg·hm-2) on growth and physiological characteristics of mulberry grown in the field using mulberry variety Longsang No.l and "qinglong" as experimental material. The results showed that leaf number, branch number, leaf area, leaf weight and leaf yield per plant increased with nitrogen application levels in0～193.2and kg·hm-20～55.2kg·hm-2, while the nitrogen application levels in241.5kg·hm-2and69kg·hm-2, the leaf area, leaf weight and leaf yield per plant reduced. Total nitrogen and total phosphorus contents in leaves of mulberry increased with nitrogen application levels in0～144.9kg·hm-2and41.4kg·hm-2, but over nitrogen application levels of that the trend is opposite, the indicating that higher nitrogen can be reduce the absorption of N and P. Chlorophyll, soluble protein contents and soluble sugar contents in leaves of mulberry increased with nitrogen application levels increased. In sum, by the N application levels on the growth and physiological characteristics of mulberry, the best N application levels in the field of "Long sang NO.1" and "qinglong" is144.9～241.5kg·hm-2and41.4～55.2kg·hm-2respectively. Nitrogen fertilizer and mulberry leaf yield amount (Y) in a four-year field experiment partial least squares regression model are as follows:
y=1552.3484111+2.702975x1+58.083982x2+9.429218x3-0.000561x1*x1-0.672614x2*x2-0.017726x3*x3-0.036371x*x2-0.005904x1*x3-0.109191x2*x3
y=1450.0240821+2.983592x1+54.401789x2+9.645707x3-0.001892x1*x1-0.579893x2*x2-0.018230x3*x3-0.032356x1*x2-0.005737x1*x3-0.102818x2*x3
y=1490.3531551+2.752968x1+51.050728x2+8.862974x3-0.001686x1*x1-0.544203x2*x2-0.016403x3*x3-0.030354x1*x2-0.005270x1*x3-0.094480x2*x3
y=608.0876801+4.590733x1+37.395394x2+5.491631x3-0.003846x1*x1-0.483353x2*x2-0.010424x3*x3-0.070032x1*x2-0.010284x1*x3-0.070982x2*x3
2. The effects of different proportions of nitrate and ammonium nitrogen applied to mulberry (Morus alba L.) on plant growth and photosynthetic characteristics of mulberry seedlings cultured in hydroponics were investigated under the same nitrogen amount using forage mulberry variety "Qinglong" as experimental material. Plant height, leaf number, leaf area and root length of mulberry seedlings grown in hydroponics of single ammonium or nitrate nitrogen source were lower than that of proportion solution of ammonium and nitrate nitrogen, leaf and root biomass of mulberry seedlings grown in hydroponics was similar trends. Plant growth and biomass in mulberry seedlings were highest while the proportions of nitrate and ammonium nitrogen were50:50and25:75. Net photosynthetic rate (Pn), stomatal conductance (Gs) and water use efficiency (WUE) in leaves of mulberry seedlings cultured in25:75proportion solution of ammonium and nitrate nitrogen were higher than that of other proportion treatments. Single ammonium or nitrate nitrogen source improved the apparent quantum efficiency (AQE) and decreased light compensation point (LCP) in leaves of mulberry seedlings. It concluded that forage mulberry varieties "Qinglong" prefer nitrate nitrogen, and the proportions of nitrate and ammonium nitrogen adapted mulberry growth were between50:50and25:75.
3. In three different nitrogen treatments, the chlorophyll content (Chl), photosynthetic
Rate (Pn), RuBPcase initial activity, RuBPcase content, mesophyll conductance (gm), carbonic Anhydrase (CA) activity, PS1activity and PS1activity of leaves were the highest in the mixture treatment of NH4++NO3-, then were in the NO3-treatment and the lowest were in NH4+treatment. No evident differences were found in RuBPcase specific activity among treatments. There was little difference on stomatol conductance (Gs) between different treatments and CK. Nitrogen nutritions decreased the intercellular CO2concentration (Ci). There were similarly Changing trends between gm, CA activity and photosynthetic rate (Pn) in different treatments, Which showed the conductivity of CO2in liquid part of mesophyll cell having probably important Influence on photosynthesis of leaves. Compared to NH4+treatment, NO3-treatment increased the photorespiration rate, the ratio of photorespiration rate to photosynthetic rate and nitrate reduetase (NR) activity in wheat leaves. The increase of photosynthetic rate in mixture treatment NH4++NO3-was mainly correlated to the higher Rubisco activity, higher photosynthetic electron conductivity and the improvement of CO2conductivity in liquid phase of mesophall cell.
4. Nitrogen plays an important role throughout the whole life of all living organisms. Nitrate and ammonium are the major forms of nitrogen available to plants. Though both forms can be utilized, great differences in plant growth are found among various plants. Since1950s many research efforts have been directed toward unraveling the causes and mechanisms of NH4+toxicity as well as the relationship of nitrate and photoprotection in plants, while present knowledge is far from complete. It is kown that most of the energy for nitrate assimilation in a cell derives from photosynthesis. Recently, nitrate reduction has been suggested to be involved in the photo protection mechanism. However, our present understanding of this role of nitrate is poor. This research focuses on the difference in allocation. The results are as follows:
1) The effects of different nitrogen forms on biomass and gas exchange of mulberry and rice were studied. Ammonium significantly decreased both shoot biomass and root biomass of mulberry, especially root biomass. Photosynthetie rate (Pn) of mulberry under ammonium condition was also significantly lower than those under nitrate condition while stomatal conductance and intracellular CO2concentration were higher. Thus, decreased Pn could be attributed to stomatal limitation. In comparison, there were no significant differences in biomass and gas exchange between rice cultured under nitrate or ammonium condition.
2) The effects of different nitrogen forms on photosynthetic electron allocation and nitrate reduetase activity were investigated in mulberry and rice plants. When cultured with ammonium as nitrogen source, mulberry plants had lower JF(PS Ⅱ)), Jo(PCR), JC(PCO. The effects of different nitrogen forms on chlorophyll fluorescence and xanthophyll cycle activity were studied in mulberry and rice plants. There was no significant difference in either chlorophyll fluorescence or xanthophyll cycle activity between nitrate and ammonium feed rice plants.
5. The responses of photosynthetic electron transport and excitation energy distribution in winter wheat leaves at the jointing stage to nitrogen levels were studied in the field experiment, both leaf gas exchange parameter and chlorophyll fluorescence were measured. The light energy absorbed by antenna pigment of both cultivars increased with the treatment of nitrogen. Although nitrogen could not obviously change the distribution proportion of photosynthetic carbon reduction and photosynthetic carbon oxidation, nitrogen could increase JF of PSII and photosynthetic rate. Nitrogen could affect PSII activities of both cultivars, and there were significant differences between the two cultivars. Nitrogen application improved the competition between heat dissipation and photo-chemic reaction and enhanced the self-protection ability of photosynthetic apparatus.
In summary, the regulation mechanism of growth of mulberry trees from nitrogen and nitrogen forms, the mulberry leaf photosynthetic electron transport and absorption of light energy distribution characteristics for clues to reveal the nitrogen, especially of nitrogen forms in Mulberry metabolism processsource of reducing power (NADPH or NADH) to clarify the reducing power of nitrate reduction to ammonium nitrogen needs from the photosynthetic electron transport chain (PS I side), but the ammonium nitrogen assimilation into amino acids in plant roots, reducing power mainly from respiration, which proved effective consumption of energy excess the photosynthetic chain electron acceptor library part of the excess excitation energy, thereby reducing the destruction of photoinhibition of photosystem and explain long beencontroversial reduction of nitrate to the contribution of the optical destruction defense mechanism, to help us further understand plant Photoprotective Mechanisms.
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