披针叶茴香生理生态学特性研究
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摘要
披针叶茴香(Illicium lanceolatum A.C.Smith)为中国所特有的传统药用植物。其所含的莽草酸(Shikimic Acid)具有较强的抗炎、镇痛和抑制血小板聚集、抑制动、静脉血栓及脑血栓形成,还可作为抗病毒和抗癌药物的中间体。这一发现使得以披针叶茴香为主的八角属植物受到前所未有的关注,其资源开发利用前景可观。披针叶茴香虽有较广的生态幅,但多处于零星分布,加之当前资源掠夺式利用方式,野生资源日渐稀少,其规模引种保护和栽培日益迫切。本研究以不同产地披针叶茴香1年生幼苗、不同自然栽培方式下4年生栽培幼苗以及1年生盆栽苗为研究对象,结合野外调查、自然栽培以及室内培养试验等研究方式,从其光合生理生态以及水分生理生态研究角度,阐明披针叶茴香栽培幼苗对生长环境生理生态响应的季节差异、产地间差异以及幼苗对不同栽培方式的适应性与生理生态响应机制,探讨1年生幼苗对光照和水分二者综合影响的响应机理,取得以下结论:
1、披针叶茴香1年生幼苗光合特性存在明显的季节差异和产地间差异。披针叶茴香净光合速率(Pn)日变化在生长初期呈“单峰型”;盛期和末期呈“双峰型”,12:00左右发生“光合午休”。披针叶茴香Pn日均值季节差异明显,表现出盛期>初期>末期的季节变化趋势。披针叶茴香WUE日变化季节性差异较明显,表现出生长末期>盛期>初期的季节变化趋势。生长初期,披针叶茴香最大净光合速率(Pmax)在2.585-5.103 umol·m~(-2)·s~(-1)范围之间;光饱和点(LSP)在373.5-1574.4 umol·m~(-2)·s~(-1)范围,光补偿点(LCP)在26.3 -61.8 umol·m~(-2)·s~(-1)范围。在生长盛期,披针叶茴香Pmax在4.6-9.9 umol·m~(-2)·s~(-1)范围变动,LSP变动范围在997.9-2182.9 umol·m~(-2)·s~(-1)之间,LCP变动范围在23.5-135.7 umol·m~(-2)·s~(-1)之间。在末期,披针叶茴香Pmax在0.151-1.163 umol·m~(-2)·s~(-1)范围变动,LSP在71.6-922.0 umol·m~(-2)·s~(-1)之间,LCP在11.1-41.3 umol·m~(-2)·s~(-1)之间。披针叶茴香光合参数季节差异表明,其Pmax、AQY以及LSP和LCP均呈生长盛期>初期>末期的季节变化趋势。净光合速率(Pn)与主要环境因子的相关分析表明,光合有效辐射(PAR)在初期、盛期不是主要制约因子,但末期却成为主要影响因子。4个产地植株对光照利用能力高低主要受季节影响,表现在LA植株生长初期和末期
对弱光利用最低、对强光利用能力较高;到了盛期,对弱光的利用能力提高。WN产地和KH产地植株初期对强光和弱光的利用能力整体水平居中;盛期和末期,WN产地对强光和弱光利用效率仍然居中,而KH产地对低光的利用能力最强。NP产地初期主要趋向于利用弱光,对强光利用能力最低;盛期则趋向于利用强光,末期弱光和强光的利用能力均处于中等水平。WN产地所表现出的对弱光和强光在全生长季节较为稳定的利用效率以及其盛期具有最高的光饱和点、末期具有较低的光补偿点的光合特性,表明WN产地对光照的适应性范围最广,揭示出WN产地是目前苗期表现较好、生态适应性最强的一个优良种源。
2、通过探讨不同栽培方式下4年生披针叶茴香植株光合生理生态特性,发现:4年生披针叶茴香植株光合日变化和个体光合特性既因栽培方式不同而不同,亦存在季节性差异;这种差异很好地体现了披针叶茴香幼苗对生长环境的光合生理适应性。光合日变化结果显示,A处理盛期较低WUE和SUE日均值表明披针叶茴香生长盛期采取遮荫措施的必要性;D处理盛期较高的SUE日均值和较低的WUE日均值体现其对弱光环境的一种适应性。不同栽培方式下披针叶茴香植株净光合速率日均值大小排序与其各生长季节环境光合有效辐射日均值大小排序密切正相关,充分体现不同栽培方式下主要小气候因子对植株光合能力的潜在影响。
光合参数特性表明,生长初期,D处理幼苗拥有最低的LCP、较高的AQY和最高的Pmax以及较高的LSP,说明其对弱光、强光的利用能力较强;A处理植株则拥有最低的LSP值、较低的LCP值和Pmax值,以及最高的AQY值,表明全光照下植株利用弱光的能力虽然较强,但在强光照下光合能力明显下降。到了生长盛期,A处理幼苗具有最低的AQY值和LSP值、较低的Pmax值以及最高的LCP值,说明此处理幼苗对强光、弱光的光合利用效率下降为最低水平,导致植株生长受限;D处理幼苗拥有最低的LCP值、较低的LSP值和较大的AQY值以及较低的Pmax值,说明D处理幼苗对弱光环境适应性最好。综合表明了D处理套种模式为披针叶茴香幼苗栽培最佳模式。
3、披针叶茴香幼苗叶绿素含量季节差异明显。叶片Chla/Chlb比值呈生长末期>盛期>初期的季节变化趋势;叶绿素总含量则呈生长盛期>初期>末期;叶绿素a和b含量分别是生长盛期>末期>初期和生长初期>盛期>末期。披针叶茴香叶绿素总含量表现出同其Pn日均值相一致的季节变化趋势。不同产地、不同栽培方式下披针叶茴香植株叶绿素含量、比叶重、叶氮含量与其光合能力的相关性受生长季节影响;4个指标大小季节性差异与其植株光合能力季节性差异密切相关。披针叶茴香幼苗形态可塑性较强,通过降低比叶重来适应弱光环境。
4、短期水分胁迫可一定程度上提高披针叶茴香的WUE和SUE值;但受水分条件影响远高于受光强影响。在低光范围(0-200umol·m~(-2)·s~(-1))时,披针叶茴香幼苗光合响应主要受土壤水分条件的影响,当环境光合有效辐射继续增强时,其光合响应受光照的负面影响大于水分影响。因此,披针叶茴香生长环境需要一定土壤水分条件,但更重要的是需要一定光照,但光强不宜太大,尤其在土壤水分条件较差时需要弱光环境来缓和逆境胁迫。WN产地植株对水分和光照的适应能力相对较强,其次为NP产地,而LA和KH产地对于水分的要求较高。
5、正常供水下,披针叶茴香幼苗生长适应光照范围变得较广。在水分胁迫下,在较低(25 umol·m~(-2)·s~(-1))或者较高光照(75 umol·m~(-2)·s~(-1))下其幼苗生长都将受限;在较高光照下,则主要受水分条件影响,充足供水下能较好的适应。无论水分条件,其比较适合50 umol·m~(-2)·s~(-1)光照下生长。披针叶幼苗整体较不耐旱,证明其喜阴湿的生物学特性。试验结果表明,正常供水下,LA产地植株比较适应在50 umol·m~(-2)·s~(-1)光照下生长;WN产地在50-75 umol·m~(-2)·s~(-1)之间较为合适;KH产地幼苗适应大于25 umol·m~(-2)·s~(-1)的光照下生长;NP产地不适应在25 umol·m~(-2)·s~(-1)以下的弱光生长,对较高光照的响应最敏感。在水分胁迫下,WN产地对光照需求范围最宽;而LA产地则最不能忍受干旱。
总之,从光合与水分生理生态角度看,WN产地是所有试验材料中生态适应性较强、对光照和水分需求性范围较宽的优良可供选择种源,其次为LA产地。80%光郁闭下套种且套种树种为生长末期落叶的光照环境较为适宜,其次为50%光郁闭下的栽培方式。
Illicium lanceolatum is a traditional and proper medicinal plant in China. The shikimic acid that is contained in I. lanceolatum has strong functions of anti-inflammatory, analgesic and inhibiting platelet aggregation, arterial and venous thrombosis and cerebral thrombosis, but also as is the intermediates for anti-virus and anti-cancer drugs. This discovery makes I. lanceolatum and other genuses of Illicium L. have been given the unprecedented attentions, and its prospects for the development and utilization are more great. Although I. lanceolatum has broader ecological amplitude, but its wild resources are scattered, the current predatory use and increasing scarcity of wild resources is more strong, which makes the protection and cultivation of its is increasingly urgent. Based on 1-year-old cultivated and potted seedlings of 4 provenances, the 4-year-old cultivations of seedlings of I. lanceolatum growthing under 5 different nature light conditions, through field and laboratory tests research, this study discusses the adaptable difference of ecophysiological responses of 4 different provenances of I. lanceolatum and LA provenance growthing under 5 natural light conditons to environment from two aspects of photosynthetic ecophysiology and water ecophysiology. The results as follows:
(1) The obvious diversities of photosynthetic characteristics of 4 provenances seedlings are existed in provenances and different growth stages. LA provenance seedlings have lowest use efficiency of low-light and more higher use efficiency of high-light during the initial and later phase, but with the higher use efficiency of low-light during prosperous phase. WN provenance has middle use efficiency for using low-light and high-light in the whole growth phase, and KH provenance has highest use efficiency for low-light during the prosperous and later phase. Whereas, NP provenance has higher use efficiency of low-light during initial phase, but of high-light during prosperous phase. The photosynthetic characteristics with highest LSPin prosperous phase and lowest LCP in later phase indicate that WN provenance is a good provenance with a widest range of adaptability to illumination. The correlation analysis between Net photosynthetic rate (Pn) and the major environmental factors show that the photosynthetic active radiation (PAR) in the early and prosperous period is not a restricted factor, but in the later period has become a major factor.
(2) The diurnal variation of photosynthesis and photosynthetic characteristics of 4-year-old I. lanceolatum seedlings under different light conditions are measured and the results show that they have obvious differences not only because of light conditions variation, there is also a seasonal differences. This difference reflects the photosynthetic physiological adaptation of seedlings to growth environment.
The lower mean diurnal value of WUE and SUE of A light treatment shows that the shading is essential during prosperous stage. The more higher mean diurnal value of SUE and more lower its of WUE embody that D light treatment is adaptable to low-light environment. The order of diurnal mean value of Pn for I. lanceolatum seedlings under 5 different light conditions is positively correlative to its of PAR.
The comparation of photosynthetic parameters reflect that D light treatment has the lowest LCP, higher AQY and LSP, highest Pmax in initial stage, which illuminates that seedlings under D environment have better light energy utilization efficiency. Whereas, it has the lowest LCP, lower LSP, lower Pmax and higher AQY in prosperous stage, which show that it is greatly adaptable to low-light environment in this stage. For A light treatment, it has lowest LSP, lower LCP and highest AQY in initial stage and has lowest AQY and LSP and highest LCP in prosperous stage, which reveal that seedlings of I. lanceolatum haven’t adaptable to entire light environment. So D light treatment is most better cultivation pattern of I. lanceolatum.
(3) The pertinency between chlorophyll content, leaf mass per area, leaf nitrogen content and photosynthetic capacity of seedlings of 4 provenance and those under 5 different light conditions is greatly affected by growing season. The seedlings of I. lanceolatum have strong morphological plasticity and are adaptable to low-light conditions by reducing the leaf mass per area.
(4) The drought stress test for 4 provenances shows that seedlings of WN provenance have strongest adaptability to water and light conditions, followed by NP provenance. Whereas, the seedlings of LA and KH provenances are more greatly restricted by water condition. The short-time drought stress can improve the value of WUE and SUE at a certain extent. In low-light range (0-200 umol·m~(-2)·s~(-1)), the photosynthetic light-response of seedlings of 4 provenances is mainly affected by soil moisture conditions, and when the photosynthetic active radiation continues to enhancing, the photosynthetic response to light will have been more negatively impacted by the light than water.
(5) The illumination cultivation and drought stress tests show that under normal water condition, the seedlings of LA provenance are growing better in a light range of 0-50 umol·m~(-2)·s~(-1), those of WN provenance are adaptable to the range of 50-75 umol·m~(-2)·s~(-1), those of KH provenance are growing better above 25 umol·m~(-2)·s~(-1), and those of NP provenance aren’t adaptable to the range of 0-25 umol·m~(-2)·s~(-1). It also shows that under drought stress, seedlings of WN provenance have widest range of lighting needs, and those of LA provenance can’t tolerate drought.
In a word, from the viewpoint of photosynthetic ecophysiology and water ecophysiology, WN provenance of I. lanceolatum is a good provenance with strongest adaptability and widest range of light and water conditions, followed by LA provenance. The other important result is that the cultivation mode of inter-planting with 80% canopy density is the most suitable mode for I. lanceolatum, followed by 50% canopy density model of inter-planting.
1、披针叶茴香1年生幼苗光合特性存在明显的季节差异和产地间差异。披针叶茴香净光合速率(Pn)日变化在生长初期呈“单峰型”;盛期和末期呈“双峰型”,12:00左右发生“光合午休”。披针叶茴香Pn日均值季节差异明显,表现出盛期>初期>末期的季节变化趋势。披针叶茴香WUE日变化季节性差异较明显,表现出生长末期>盛期>初期的季节变化趋势。生长初期,披针叶茴香最大净光合速率(Pmax)在2.585-5.103 umol·m~(-2)·s~(-1)范围之间;光饱和点(LSP)在373.5-1574.4 umol·m~(-2)·s~(-1)范围,光补偿点(LCP)在26.3 -61.8 umol·m~(-2)·s~(-1)范围。在生长盛期,披针叶茴香Pmax在4.6-9.9 umol·m~(-2)·s~(-1)范围变动,LSP变动范围在997.9-2182.9 umol·m~(-2)·s~(-1)之间,LCP变动范围在23.5-135.7 umol·m~(-2)·s~(-1)之间。在末期,披针叶茴香Pmax在0.151-1.163 umol·m~(-2)·s~(-1)范围变动,LSP在71.6-922.0 umol·m~(-2)·s~(-1)之间,LCP在11.1-41.3 umol·m~(-2)·s~(-1)之间。披针叶茴香光合参数季节差异表明,其Pmax、AQY以及LSP和LCP均呈生长盛期>初期>末期的季节变化趋势。净光合速率(Pn)与主要环境因子的相关分析表明,光合有效辐射(PAR)在初期、盛期不是主要制约因子,但末期却成为主要影响因子。4个产地植株对光照利用能力高低主要受季节影响,表现在LA植株生长初期和末期
对弱光利用最低、对强光利用能力较高;到了盛期,对弱光的利用能力提高。WN产地和KH产地植株初期对强光和弱光的利用能力整体水平居中;盛期和末期,WN产地对强光和弱光利用效率仍然居中,而KH产地对低光的利用能力最强。NP产地初期主要趋向于利用弱光,对强光利用能力最低;盛期则趋向于利用强光,末期弱光和强光的利用能力均处于中等水平。WN产地所表现出的对弱光和强光在全生长季节较为稳定的利用效率以及其盛期具有最高的光饱和点、末期具有较低的光补偿点的光合特性,表明WN产地对光照的适应性范围最广,揭示出WN产地是目前苗期表现较好、生态适应性最强的一个优良种源。
2、通过探讨不同栽培方式下4年生披针叶茴香植株光合生理生态特性,发现:4年生披针叶茴香植株光合日变化和个体光合特性既因栽培方式不同而不同,亦存在季节性差异;这种差异很好地体现了披针叶茴香幼苗对生长环境的光合生理适应性。光合日变化结果显示,A处理盛期较低WUE和SUE日均值表明披针叶茴香生长盛期采取遮荫措施的必要性;D处理盛期较高的SUE日均值和较低的WUE日均值体现其对弱光环境的一种适应性。不同栽培方式下披针叶茴香植株净光合速率日均值大小排序与其各生长季节环境光合有效辐射日均值大小排序密切正相关,充分体现不同栽培方式下主要小气候因子对植株光合能力的潜在影响。
光合参数特性表明,生长初期,D处理幼苗拥有最低的LCP、较高的AQY和最高的Pmax以及较高的LSP,说明其对弱光、强光的利用能力较强;A处理植株则拥有最低的LSP值、较低的LCP值和Pmax值,以及最高的AQY值,表明全光照下植株利用弱光的能力虽然较强,但在强光照下光合能力明显下降。到了生长盛期,A处理幼苗具有最低的AQY值和LSP值、较低的Pmax值以及最高的LCP值,说明此处理幼苗对强光、弱光的光合利用效率下降为最低水平,导致植株生长受限;D处理幼苗拥有最低的LCP值、较低的LSP值和较大的AQY值以及较低的Pmax值,说明D处理幼苗对弱光环境适应性最好。综合表明了D处理套种模式为披针叶茴香幼苗栽培最佳模式。
3、披针叶茴香幼苗叶绿素含量季节差异明显。叶片Chla/Chlb比值呈生长末期>盛期>初期的季节变化趋势;叶绿素总含量则呈生长盛期>初期>末期;叶绿素a和b含量分别是生长盛期>末期>初期和生长初期>盛期>末期。披针叶茴香叶绿素总含量表现出同其Pn日均值相一致的季节变化趋势。不同产地、不同栽培方式下披针叶茴香植株叶绿素含量、比叶重、叶氮含量与其光合能力的相关性受生长季节影响;4个指标大小季节性差异与其植株光合能力季节性差异密切相关。披针叶茴香幼苗形态可塑性较强,通过降低比叶重来适应弱光环境。
4、短期水分胁迫可一定程度上提高披针叶茴香的WUE和SUE值;但受水分条件影响远高于受光强影响。在低光范围(0-200umol·m~(-2)·s~(-1))时,披针叶茴香幼苗光合响应主要受土壤水分条件的影响,当环境光合有效辐射继续增强时,其光合响应受光照的负面影响大于水分影响。因此,披针叶茴香生长环境需要一定土壤水分条件,但更重要的是需要一定光照,但光强不宜太大,尤其在土壤水分条件较差时需要弱光环境来缓和逆境胁迫。WN产地植株对水分和光照的适应能力相对较强,其次为NP产地,而LA和KH产地对于水分的要求较高。
5、正常供水下,披针叶茴香幼苗生长适应光照范围变得较广。在水分胁迫下,在较低(25 umol·m~(-2)·s~(-1))或者较高光照(75 umol·m~(-2)·s~(-1))下其幼苗生长都将受限;在较高光照下,则主要受水分条件影响,充足供水下能较好的适应。无论水分条件,其比较适合50 umol·m~(-2)·s~(-1)光照下生长。披针叶幼苗整体较不耐旱,证明其喜阴湿的生物学特性。试验结果表明,正常供水下,LA产地植株比较适应在50 umol·m~(-2)·s~(-1)光照下生长;WN产地在50-75 umol·m~(-2)·s~(-1)之间较为合适;KH产地幼苗适应大于25 umol·m~(-2)·s~(-1)的光照下生长;NP产地不适应在25 umol·m~(-2)·s~(-1)以下的弱光生长,对较高光照的响应最敏感。在水分胁迫下,WN产地对光照需求范围最宽;而LA产地则最不能忍受干旱。
总之,从光合与水分生理生态角度看,WN产地是所有试验材料中生态适应性较强、对光照和水分需求性范围较宽的优良可供选择种源,其次为LA产地。80%光郁闭下套种且套种树种为生长末期落叶的光照环境较为适宜,其次为50%光郁闭下的栽培方式。
Illicium lanceolatum is a traditional and proper medicinal plant in China. The shikimic acid that is contained in I. lanceolatum has strong functions of anti-inflammatory, analgesic and inhibiting platelet aggregation, arterial and venous thrombosis and cerebral thrombosis, but also as is the intermediates for anti-virus and anti-cancer drugs. This discovery makes I. lanceolatum and other genuses of Illicium L. have been given the unprecedented attentions, and its prospects for the development and utilization are more great. Although I. lanceolatum has broader ecological amplitude, but its wild resources are scattered, the current predatory use and increasing scarcity of wild resources is more strong, which makes the protection and cultivation of its is increasingly urgent. Based on 1-year-old cultivated and potted seedlings of 4 provenances, the 4-year-old cultivations of seedlings of I. lanceolatum growthing under 5 different nature light conditions, through field and laboratory tests research, this study discusses the adaptable difference of ecophysiological responses of 4 different provenances of I. lanceolatum and LA provenance growthing under 5 natural light conditons to environment from two aspects of photosynthetic ecophysiology and water ecophysiology. The results as follows:
(1) The obvious diversities of photosynthetic characteristics of 4 provenances seedlings are existed in provenances and different growth stages. LA provenance seedlings have lowest use efficiency of low-light and more higher use efficiency of high-light during the initial and later phase, but with the higher use efficiency of low-light during prosperous phase. WN provenance has middle use efficiency for using low-light and high-light in the whole growth phase, and KH provenance has highest use efficiency for low-light during the prosperous and later phase. Whereas, NP provenance has higher use efficiency of low-light during initial phase, but of high-light during prosperous phase. The photosynthetic characteristics with highest LSPin prosperous phase and lowest LCP in later phase indicate that WN provenance is a good provenance with a widest range of adaptability to illumination. The correlation analysis between Net photosynthetic rate (Pn) and the major environmental factors show that the photosynthetic active radiation (PAR) in the early and prosperous period is not a restricted factor, but in the later period has become a major factor.
(2) The diurnal variation of photosynthesis and photosynthetic characteristics of 4-year-old I. lanceolatum seedlings under different light conditions are measured and the results show that they have obvious differences not only because of light conditions variation, there is also a seasonal differences. This difference reflects the photosynthetic physiological adaptation of seedlings to growth environment.
The lower mean diurnal value of WUE and SUE of A light treatment shows that the shading is essential during prosperous stage. The more higher mean diurnal value of SUE and more lower its of WUE embody that D light treatment is adaptable to low-light environment. The order of diurnal mean value of Pn for I. lanceolatum seedlings under 5 different light conditions is positively correlative to its of PAR.
The comparation of photosynthetic parameters reflect that D light treatment has the lowest LCP, higher AQY and LSP, highest Pmax in initial stage, which illuminates that seedlings under D environment have better light energy utilization efficiency. Whereas, it has the lowest LCP, lower LSP, lower Pmax and higher AQY in prosperous stage, which show that it is greatly adaptable to low-light environment in this stage. For A light treatment, it has lowest LSP, lower LCP and highest AQY in initial stage and has lowest AQY and LSP and highest LCP in prosperous stage, which reveal that seedlings of I. lanceolatum haven’t adaptable to entire light environment. So D light treatment is most better cultivation pattern of I. lanceolatum.
(3) The pertinency between chlorophyll content, leaf mass per area, leaf nitrogen content and photosynthetic capacity of seedlings of 4 provenance and those under 5 different light conditions is greatly affected by growing season. The seedlings of I. lanceolatum have strong morphological plasticity and are adaptable to low-light conditions by reducing the leaf mass per area.
(4) The drought stress test for 4 provenances shows that seedlings of WN provenance have strongest adaptability to water and light conditions, followed by NP provenance. Whereas, the seedlings of LA and KH provenances are more greatly restricted by water condition. The short-time drought stress can improve the value of WUE and SUE at a certain extent. In low-light range (0-200 umol·m~(-2)·s~(-1)), the photosynthetic light-response of seedlings of 4 provenances is mainly affected by soil moisture conditions, and when the photosynthetic active radiation continues to enhancing, the photosynthetic response to light will have been more negatively impacted by the light than water.
(5) The illumination cultivation and drought stress tests show that under normal water condition, the seedlings of LA provenance are growing better in a light range of 0-50 umol·m~(-2)·s~(-1), those of WN provenance are adaptable to the range of 50-75 umol·m~(-2)·s~(-1), those of KH provenance are growing better above 25 umol·m~(-2)·s~(-1), and those of NP provenance aren’t adaptable to the range of 0-25 umol·m~(-2)·s~(-1). It also shows that under drought stress, seedlings of WN provenance have widest range of lighting needs, and those of LA provenance can’t tolerate drought.
In a word, from the viewpoint of photosynthetic ecophysiology and water ecophysiology, WN provenance of I. lanceolatum is a good provenance with strongest adaptability and widest range of light and water conditions, followed by LA provenance. The other important result is that the cultivation mode of inter-planting with 80% canopy density is the most suitable mode for I. lanceolatum, followed by 50% canopy density model of inter-planting.
引文
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