油茶耐弱光生理特性研究
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摘要
油茶泛指山茶属(Camellia)植物中含油率较高的各油茶物种,在我国有悠久的栽培历史,目前对油茶的研究集中在育种、栽培等方面,在国内外对油茶光合生理的研究不多。本文在对普通油茶(C. oleifera)、广宁红花油茶(C.semiserrata)、博白大果油茶(C.gigantocarpa)、日本红山茶(C.japonica)、南荣油茶(C.nanyongensis)、香花油茶(C.osmantha Ye CX)等6个油茶物种和普通油茶中选育的岑软2号、岑软3号、岑软24号、湘林86号、长林4号、赣190号6个无性系光合特性研究的基础上,通过覆盖遮荫材料对不同强度(60%、75%、90%)遮荫处理下5个油茶物种(普通油茶、广宁红花油茶、博白大果油茶、南荣油茶,香花油茶)和5个油茶优良无性系(岑软2号、岑软3号、岑软24号、湘林86号、赣190号)苗期光合及生长特性进行调查,并通过模拟高大乔木下的自然荫蔽环境对广西(岑软3号)、湖南(湘林86号)和江西(赣190号)选育的优良无性系成年植株在弱光胁迫下的光合及生长发育特性进行研究,旨在为油茶复层栽培提供理论依据。研究结果如下：
(1)不同油茶物种、普通油茶不同无性系光合特性存在差异。不同油茶物种日平均净光合速率由大到小依次为普通油茶>香花油茶>博白大果油茶>广宁红花油茶>日本红山茶>宛田红花油茶,其中日平均净光合速率最高的普通油茶达到7.24μmol·m-2·s-1,而最低的宛田红花油茶日平均净光合速率为3.73μmol·m-2·s-1,仅为普通油茶的51.52%。参试普通油茶不同无性系日平均净光合速率最大为湘林86号,达到6.86μmol·m-2·s-1,最小的岑软2号仅为3.97μmol·m-2·s-1,是湘林86号的57.87%。日平均净光合速率由大到小依次排列为湘林86号>岑软24号>长林4号>岑软3号>赣190号>岑软2号。
(2)在模拟光辐射条件下,不同油茶物种最大净光合速率在8.4～10.5μmol·m-2·s-1之间,其中普通油茶最高,宛田红花油茶最低；普通油茶的光补偿点仅为15.50μmol·m-2·s-1,利用弱光的能力强,其次是香花油茶和日本红山茶,宛田红花油茶、广宁红花油茶和博白大果油茶利用弱光能力较差；分布在低纬度地区的香花油茶光饱和点最高,最耐强光,分布在高纬度地区的日本红山茶光饱和点最低,利用强光的能力最差。参试油茶无性系中,最大净光合速率在5.8～11.4μmol·m-2·s-1之间,岑软3号最高,岑软2号最低；赣190号光补偿点为16.36μmol·m-2·s-1,利用弱光能力最强,岑软2号和长林4号利用弱光能力较差；岑软3号光饱和点为537.86μmol·m-2·s-1,利用强光的能力最强,赣190号利用强光的能力最差,光饱和点仅为383.55μmol·m-2·s-1。
(3)9月底、10月初,未遮荫条件下的参试油茶物种苗期、普通油茶无性系苗期和成年期净光合速率日变化曲线有单峰型和双峰型,其中油茶物种博白大果油茶苗期,普通油茶无性系岑软24号苗期以及湘林86号成年期为单峰曲线,剩余油茶物种普通油茶、香花油茶、广宁红花油茶和宛田红花油茶苗期,普通油茶无性系岑软2号、岑软3号、湘林86号、赣190号苗期,以及普通油茶无性系岑软3号、赣190号成年期均为双峰曲线。但随着遮荫强度的增大,所有参试油茶物种苗期、普通油茶无性系苗期和成年期净光合速率日变化曲线均为单峰型。在60%-90%遮荫条件下所有参试油茶物种苗期、普通油茶无性系苗期和成年期日平均净光合速率均低于对照,但日平均净光合速率与对照的比值都大于相对应遮荫条件下光合有效辐射与对照的比值,表明在60%～90%遮荫条件下参试的所有油茶物种、普通油茶无性系叶片捕捉弱光进行光合作用的能力增强。
(4)油茶物种广宁红花油茶、普通油茶、宛田红花油茶、香花油茶苗期,普通油茶无性系岑软2号、岑软3号、湘林86号苗期,以及普通油茶无性系岑软3号、湘林86号、赣190号成年期在60%-90%遮荫下叶片叶绿素a、叶绿素b和类胡萝卜素含量都显著增加；油茶物种博白大果油茶以及普通油茶无性系岑软24号和赣190号在75%～90%遮荫下叶片叶绿素a、叶绿素b和类胡萝卜素也显著高于对照。弱光下促进了叶片叶绿素和类胡萝卜素的合成与其在遮荫条件下利用弱光进行光合作用的能力增强的变化趋势相一致。在弱光下,参试油茶物种以及普通油茶无性系苗期Qchlb/Qchla增大幅度不明显,叶绿素合成过程中并没有优先合成叶绿素b,但除油茶物种普通油茶外,剩余油茶物种以及普通油茶无性系Qchl/Qcar都显著高于对照,叶绿素增大幅度都要大于类胡萝卜素,叶片色素合成中优先合成了叶绿素。而普通油茶无性系成年期中,除90%遮荫条件下的赣190号Qchl/Qcar显著高于对照外,其余遮荫处理Qchl/Qcar没有显著增大,优先合成叶绿素的趋势不明显,而75%和90%遮荫处理的3个参试油茶无性系以及60%遮荫处理的湘林86号和赣190号叶片Qchlb/Qchla明显增大,叶绿素合成中优先合成了叶绿素b,从而更好的捕捉散射光。
(5)苗期,参试油茶物种中博白大果油茶和广宁红花油茶耐弱光能力最强,90%遮荫下不仅对其根、茎和叶片的生长没有抑制,且60%强度遮荫下促进了根系的生长；普通油茶和香花油茶也有较强的耐弱光能力,各遮荫处理对其根、茎和叶片的生长都没有影响,宛田红花油茶苗期相对耐弱光能力较差,60%遮荫处理下的宛田红花油茶根系生长受到明显抑制。参试普通油茶不同油茶无性系中,岑软2号和赣190号苗期耐弱光能力最强,各遮荫处理不仅对其根、茎、叶的生长没有抑制作用,且部分强度遮荫处理促进了植株的营养生长；岑软24号耐弱光能力也较强,虽然90%遮荫条件下根系生长受到抑制,但部分遮荫处理却却促进了茎和叶片的生长；岑软3号和湘林86号苗期耐弱光的能力相对较差,75%遮荫处理会明显抑制根系的生长。
(6)对参试普通油茶无性系成年期植株遮荫22个月后调查表明,岑软3号和湘林86号在90%遮荫条件下以及赣190号在60%～90%遮荫条件下枝条的生长都朝着更有利于捕捉光的方向发展,同时,90%遮荫处理还促进了湘林86号和赣190号叶片朝着捕捉光的方向生长,但各遮荫处理下的3个无性系叶片N、P、K元素的含量与对照都没有显著差异。在60%遮荫条件下,岑软3号能正常花芽分化、座果,且果实质量和油脂组分与对照都没有显著差异,赣190号虽然花芽数量、座果数量以及油脂组分与对照都没有显著性差异,但果实膨大却受到抑制,而湘林86号花芽分化和座果都显著低于对照；虽然,75%和90%遮荫处理后的岑软3号、湘林86号和赣190号果实油脂组分与对照差异不显著,但花芽分化和座果都受到明显抑制。成年期,普通油茶3个无性系耐弱光能力由大到小依次为岑软3号>赣190号>湘林86号。
(7)苗期试验表明,不同油茶物种以及普通油茶无性系都具有较强的耐弱光能力,博白大果油茶、广宁红花油茶、普通油茶、香花油茶,以及普通油茶无性系岑软2号和赣190号可以在90%以下遮荫(10月初日平均光合有效辐射为61.83μmol·m-2·s-1)条件下正常生长；岑软24号在75%(10月初日平均光合有效辐射为178.87μmol·m-2·s-1)以下遮荫条件下能正常生长；而岑软3号和湘林86号也能在60%(10月初日平均光合有效辐射为311.91μmol·m-2·s-1)以下遮荫条件下生长。普通油茶无性系成年期耐弱光能力相对较弱,75%(9月30日日平均光合有效辐射为193.25μmol·m-2·s-1)以上遮荫条件下岑软3号、湘林86号和赣190号花芽分化和座果受到抑制,但岑软3号在60%(9月30日平均光合有效辐射为280.72μmol·m-2·s-1)遮荫条件下能正常生长、发育,赣190号也能正常开花,座果。
With a long cultivation history in China, oil-tea camellia is a general reference to a section of species that are relatively high in oil content in genus Camellia. Research on Camellia was mainly concentrated in the fields of conventional breeding and cultivation measures for a long time, yet few studies were conducted on photosynthetic physiology in China and overseas. For the purpose of providing theoretical basis for compound cultivation, based on the photosynthetic properties of six camellia species (C. oleifera, C. vietnamensis, C. gigantocarpa, C. japonica, C. semiserrata, and C. osmantha) and six clones selected from C. oleifera (Cenruan2, Cenruan3, Cenruan24, Xianglin86, Changlin4and Gan190), investigation was carried out on photosynthetic and growth properties during seedling stage of five camellia species (C. oleifera, C. vietnamensis, C. gigantocarpa, C. semiserrata, C. osmantha) and five superior clones (Cenruan2, Cenruan3, Cenruan24, Xianglin86and Gan190) under three levels of light intensity treatments (60%,75%and90%), and studies were done on photosynthetic and growth properties of selected superior clones from Guangxi (Cenruan3), Hunan (Xianglin86) and Jiangxi (Gan190) at adult stage under the condition of low-light stress by simulating a natural shading environment under high arbor trees. Research results are as follows:
(1) There were differences in photosynthetic properties among different camellia species and different clones of C. oleifera. The order of daily average net photosynthetic efficiency of different camellia species was C. oleifera>C. osmantha-> C. gigantocarpa> C. vietnamensis> C. japonica> C. polyodonta. The daily average net photosynthetic efficiency of C.oleifera was the highest, at7.24μmol·m-2·s-1, and C.polyodonta was the lowest, at3.72μmol·m-2·s-1, merely taking up51.52%of C. oleifera. Among the tested C. oleifera clones, photosynthetic properties of Xianglin86was the highest, at6.86μmol·m-2·s-1, Cenruan2was the lowest at3.97μmol·m-2·s-1, only taking up57.87%of Xianglin86. The daily average net photosynthetic efficiency followed this order:Xianglin86> Cenruan24> Changlin4> Cenruan3> Gan190> Cenruan2, respectively.
(2) By simulating the light radiation condition, the maximum net photosynthetic efficiency of different camellia species ranged between8.4and10.5μmol·m-2·s-1, with C.oleifera the highest and C. polyodonta the lowest. Light compensation point of C.oleifera was merely15.50μmol·m-2·s-1, with the high capacity to utilize low light, followed by C. osmantha and C.japonica. C. polyodonta, C. vietnamensis and C. gigantocarpa had weak capacity to utilize low light. Light saturation point of C. osmantha which was distributed in areas of low latitude, showed high tolerance against strong light, while C.japonica which was distributed at high latitude showed the opposite. Among the tested clones, the maximum net photosynthetic efficiency ranged between5.8and11.4μmol·m-2·s-1, Cenruan3the highest and Cenruan2the lowest. The light compensation point of Gan190was16.36μmol·m-2·s-1,with the strongest capacity to utilize low light, while Cenruan2and Changlin4showed the weak capacity to utilize low light. Light saturation point of Cenruan3was537.86μmol·m-2·s-1, with the highest capacity to utilize strong light, while Gan190had the weakest capacity to utilize low light, with the light saturation point at383.55μmol·m-2·s-1.
(3) In late September and early October, under the shading-free condition, the curve of diurnal variation of net photosynthetic rate of the tested camellia species during seedling stage as well as C. oleifera clones during seedling and adult stage demonstrated single and double apexes. C. gigantocarpa of camellia species and Cenruan24of C. oleifera clone during seedling stage and Xianglin86during adult stage showed single apexes. The remaining camellia species, Sect. Paracamellia, C. vietnamensis and C. polyodonta during the seedling stage, and Cenruan2, Cenruan3, Xianglin86and Gan190of C. oleifera clones during the seedling stage, as well as Cenruan3and Gan190during the adult stage showed double apexes. Along with the increase of shading intensity, the curve of diurnal variation of net photosynthetic rate of all the camellia species during seedling stage and C. oleifera clones during seedling and adult stage showed single apexes. Under the condition of60%-90%shading, the curve of diurnal variation of net photosynthetic rate of all tested camellia species during seedling stage and C. oleifera clones during seedling and adult stage was lower than the control. Ratio of daily average net photosynthetic efficiency against that of control was higher than the ratio of photosynthetic active radiation against that of control under the same shading condition, indicating that under the condition of60%-90%shading, capacity of leaves of all tested camellia species and C.oleifera clones to capture low light was increased.
(4) Content of chlorophyll a, b and carotenoid in leaves of C. vietnamensis, C. oleifera, C. polyodonta, C. osmantha, Cenruan2, Cenruan3and Xianglin86during seedling stage and Cenruan3, Xianglin86and Gan190during adult stage under60%-90%shading condition increased obviously. Content of chlorophyll a, b and carotenoid in leaves of C. gigantocarpa, Cenruan24and Gan190under75%-90%shading condition was obviously higher than that of control. The synthesis of leaves chlorophyll and carotenoid under low light was enhanced in consistency with the increased capacity to utilize low light under shading condition for photosynthesis. Under low-light condition, increase of Qchlb/Qchla in the tested camellia species and C. oleifera clones during seedling stage was not obvious. During the process of chlorophyll synthesis, chlorophyll b was not synthesized in priority. Except C. oleifera, Qchlb/Qchla of remaining camellia species and C. oleifera was obviously higher than that of control, the increase extent of chlorophyll was higher than that of carotenoid, and chlorophyll was synthesized in priority. Among the C. oleifera clones during adult stage, except Qchlb/Qchla of Gan190under90%shading condition was obviously higher than that of control, Qchlb/Qchla of the remaining shading treatments was not increased significantly, and tendency to synthesize chlorophyll in priority was not obvious. However, Qchlb/Qchla in leaves of three tested clones under75%and90%shading treatments as well as Xianglin86and Gan190under60%shading treatment was increased significantly, chlorophyll b was synthesized in priority to better capture scattered light.
(5) During seedling stage, among the tested camellia species, the tolerance ability against low light of C. gigantocarpa and C. vietnamensis was the highest that there was no inhibiting to the growth of root, stem and leaves under90%shading treatment condition, and the growth of root was enhanced under the situation of60%shading intensity. C. oleifera and C. osmantha had relatively strong ability of low-light tolerance; there was no significant impact of each shading treatment on the growth of root, stem and leaves. C.polyodonta had relatively weak ability of low-light tolerance during seedling stage, and the growth of root system was inhibited obviously under60%shading treatment. Among the different clones of C. oleifera, Cenruan2and Gan190had the strongest ability of low-light tolerance, there was no inhibiting effects of each treatment on the growth of root, stem and leaves, and partial shading treatment enhanced the vegetative growth of the plant. Cenruan24had a relatively strong ability of low-light tolerance, although root growth was inhibited under90%shading condition, the growth of stem and leaves was enhanced under partial shading condition. Cenruan3and Xianglin86during seedling stage had a relatively weak ability of low-light tolerance, and75%shading would obviously inhibit the growth of root.
(6) Twenty-two months after shading treatment was applied on the tested C. oleifera clones during adult stage, it was found out that growth of Cenruan3and Xianglin86under90%shading condition and Gan190under60%-90%shading condition was prone to capturing light. Meanwhile,90%shading treatment also enhanced the growth of Xianglin86and Gan190towards capturing light. However, there was no significant difference in the content of N, P and K between three clones and control under each shading treatment. Under60%shading condition, Cenruan3had normal bud initiation and fruits, and there was no significant difference in fruit quality and oil composition compared with control. Though Gan190had similar bud quantity, fruit quantity and oil composition compared with control, fruit enlargement was inhibited. Bud initiation and fruits of Xianglin86were significantly lower than that of control. Although oil composition of fruits of Cenruan3, Xianglin86and Gan190under75%and90%shading condition was similar with that of control, bud initiation and fruits were obviously inhibited. At the adult stage, ability of low-light tolerance of three clones followed this order:Cenruan3>Gan190>Xianglin86.
(7) At seedling stage, camellia species and C. oleifera clones all had relatively strong ability of low-light tolerance. C. gigantocarpa, C. vietnamensis, C. oleifera, C. osmantha, Cenruan2and Gan190of C. oleifera could grow normally under90%shading condition (in early October photosynthetic active radiation was61.83μmol·m-2·s-1). Cenruan24could grow normally under75%shading condition (in early October photosynthetic active radiation was178.87μmol·m-2·s-1). Cenruan3and Xianglin86could also grow normally under60%shading condition (in early October photosynthetic active radiation was311.91μmol·m-2·s-1). C. oleifera clones at adult stage had relatively weak ability of low-light tolerance. Bud initiation and fruits of Cenruan3, Xianglin86and Gan190under75%shading condition (on September30th the diurnal average synthetic active radiation was193.25μmol·m-2·s-1) were inhibited. However, Cenruan3could grow normally under60%shading condition (on September30th the diurnal average synthetic active radiation was280.72μmol·m-2·s-1), and Gan190could also flower and have fruits normally.
(1)不同油茶物种、普通油茶不同无性系光合特性存在差异。不同油茶物种日平均净光合速率由大到小依次为普通油茶>香花油茶>博白大果油茶>广宁红花油茶>日本红山茶>宛田红花油茶,其中日平均净光合速率最高的普通油茶达到7.24μmol·m-2·s-1,而最低的宛田红花油茶日平均净光合速率为3.73μmol·m-2·s-1,仅为普通油茶的51.52%。参试普通油茶不同无性系日平均净光合速率最大为湘林86号,达到6.86μmol·m-2·s-1,最小的岑软2号仅为3.97μmol·m-2·s-1,是湘林86号的57.87%。日平均净光合速率由大到小依次排列为湘林86号>岑软24号>长林4号>岑软3号>赣190号>岑软2号。
(2)在模拟光辐射条件下,不同油茶物种最大净光合速率在8.4～10.5μmol·m-2·s-1之间,其中普通油茶最高,宛田红花油茶最低；普通油茶的光补偿点仅为15.50μmol·m-2·s-1,利用弱光的能力强,其次是香花油茶和日本红山茶,宛田红花油茶、广宁红花油茶和博白大果油茶利用弱光能力较差；分布在低纬度地区的香花油茶光饱和点最高,最耐强光,分布在高纬度地区的日本红山茶光饱和点最低,利用强光的能力最差。参试油茶无性系中,最大净光合速率在5.8～11.4μmol·m-2·s-1之间,岑软3号最高,岑软2号最低；赣190号光补偿点为16.36μmol·m-2·s-1,利用弱光能力最强,岑软2号和长林4号利用弱光能力较差；岑软3号光饱和点为537.86μmol·m-2·s-1,利用强光的能力最强,赣190号利用强光的能力最差,光饱和点仅为383.55μmol·m-2·s-1。
(3)9月底、10月初,未遮荫条件下的参试油茶物种苗期、普通油茶无性系苗期和成年期净光合速率日变化曲线有单峰型和双峰型,其中油茶物种博白大果油茶苗期,普通油茶无性系岑软24号苗期以及湘林86号成年期为单峰曲线,剩余油茶物种普通油茶、香花油茶、广宁红花油茶和宛田红花油茶苗期,普通油茶无性系岑软2号、岑软3号、湘林86号、赣190号苗期,以及普通油茶无性系岑软3号、赣190号成年期均为双峰曲线。但随着遮荫强度的增大,所有参试油茶物种苗期、普通油茶无性系苗期和成年期净光合速率日变化曲线均为单峰型。在60%-90%遮荫条件下所有参试油茶物种苗期、普通油茶无性系苗期和成年期日平均净光合速率均低于对照,但日平均净光合速率与对照的比值都大于相对应遮荫条件下光合有效辐射与对照的比值,表明在60%～90%遮荫条件下参试的所有油茶物种、普通油茶无性系叶片捕捉弱光进行光合作用的能力增强。
(4)油茶物种广宁红花油茶、普通油茶、宛田红花油茶、香花油茶苗期,普通油茶无性系岑软2号、岑软3号、湘林86号苗期,以及普通油茶无性系岑软3号、湘林86号、赣190号成年期在60%-90%遮荫下叶片叶绿素a、叶绿素b和类胡萝卜素含量都显著增加；油茶物种博白大果油茶以及普通油茶无性系岑软24号和赣190号在75%～90%遮荫下叶片叶绿素a、叶绿素b和类胡萝卜素也显著高于对照。弱光下促进了叶片叶绿素和类胡萝卜素的合成与其在遮荫条件下利用弱光进行光合作用的能力增强的变化趋势相一致。在弱光下,参试油茶物种以及普通油茶无性系苗期Qchlb/Qchla增大幅度不明显,叶绿素合成过程中并没有优先合成叶绿素b,但除油茶物种普通油茶外,剩余油茶物种以及普通油茶无性系Qchl/Qcar都显著高于对照,叶绿素增大幅度都要大于类胡萝卜素,叶片色素合成中优先合成了叶绿素。而普通油茶无性系成年期中,除90%遮荫条件下的赣190号Qchl/Qcar显著高于对照外,其余遮荫处理Qchl/Qcar没有显著增大,优先合成叶绿素的趋势不明显,而75%和90%遮荫处理的3个参试油茶无性系以及60%遮荫处理的湘林86号和赣190号叶片Qchlb/Qchla明显增大,叶绿素合成中优先合成了叶绿素b,从而更好的捕捉散射光。
(5)苗期,参试油茶物种中博白大果油茶和广宁红花油茶耐弱光能力最强,90%遮荫下不仅对其根、茎和叶片的生长没有抑制,且60%强度遮荫下促进了根系的生长；普通油茶和香花油茶也有较强的耐弱光能力,各遮荫处理对其根、茎和叶片的生长都没有影响,宛田红花油茶苗期相对耐弱光能力较差,60%遮荫处理下的宛田红花油茶根系生长受到明显抑制。参试普通油茶不同油茶无性系中,岑软2号和赣190号苗期耐弱光能力最强,各遮荫处理不仅对其根、茎、叶的生长没有抑制作用,且部分强度遮荫处理促进了植株的营养生长；岑软24号耐弱光能力也较强,虽然90%遮荫条件下根系生长受到抑制,但部分遮荫处理却却促进了茎和叶片的生长；岑软3号和湘林86号苗期耐弱光的能力相对较差,75%遮荫处理会明显抑制根系的生长。
(6)对参试普通油茶无性系成年期植株遮荫22个月后调查表明,岑软3号和湘林86号在90%遮荫条件下以及赣190号在60%～90%遮荫条件下枝条的生长都朝着更有利于捕捉光的方向发展,同时,90%遮荫处理还促进了湘林86号和赣190号叶片朝着捕捉光的方向生长,但各遮荫处理下的3个无性系叶片N、P、K元素的含量与对照都没有显著差异。在60%遮荫条件下,岑软3号能正常花芽分化、座果,且果实质量和油脂组分与对照都没有显著差异,赣190号虽然花芽数量、座果数量以及油脂组分与对照都没有显著性差异,但果实膨大却受到抑制,而湘林86号花芽分化和座果都显著低于对照；虽然,75%和90%遮荫处理后的岑软3号、湘林86号和赣190号果实油脂组分与对照差异不显著,但花芽分化和座果都受到明显抑制。成年期,普通油茶3个无性系耐弱光能力由大到小依次为岑软3号>赣190号>湘林86号。
(7)苗期试验表明,不同油茶物种以及普通油茶无性系都具有较强的耐弱光能力,博白大果油茶、广宁红花油茶、普通油茶、香花油茶,以及普通油茶无性系岑软2号和赣190号可以在90%以下遮荫(10月初日平均光合有效辐射为61.83μmol·m-2·s-1)条件下正常生长；岑软24号在75%(10月初日平均光合有效辐射为178.87μmol·m-2·s-1)以下遮荫条件下能正常生长；而岑软3号和湘林86号也能在60%(10月初日平均光合有效辐射为311.91μmol·m-2·s-1)以下遮荫条件下生长。普通油茶无性系成年期耐弱光能力相对较弱,75%(9月30日日平均光合有效辐射为193.25μmol·m-2·s-1)以上遮荫条件下岑软3号、湘林86号和赣190号花芽分化和座果受到抑制,但岑软3号在60%(9月30日平均光合有效辐射为280.72μmol·m-2·s-1)遮荫条件下能正常生长、发育,赣190号也能正常开花,座果。
With a long cultivation history in China, oil-tea camellia is a general reference to a section of species that are relatively high in oil content in genus Camellia. Research on Camellia was mainly concentrated in the fields of conventional breeding and cultivation measures for a long time, yet few studies were conducted on photosynthetic physiology in China and overseas. For the purpose of providing theoretical basis for compound cultivation, based on the photosynthetic properties of six camellia species (C. oleifera, C. vietnamensis, C. gigantocarpa, C. japonica, C. semiserrata, and C. osmantha) and six clones selected from C. oleifera (Cenruan2, Cenruan3, Cenruan24, Xianglin86, Changlin4and Gan190), investigation was carried out on photosynthetic and growth properties during seedling stage of five camellia species (C. oleifera, C. vietnamensis, C. gigantocarpa, C. semiserrata, C. osmantha) and five superior clones (Cenruan2, Cenruan3, Cenruan24, Xianglin86and Gan190) under three levels of light intensity treatments (60%,75%and90%), and studies were done on photosynthetic and growth properties of selected superior clones from Guangxi (Cenruan3), Hunan (Xianglin86) and Jiangxi (Gan190) at adult stage under the condition of low-light stress by simulating a natural shading environment under high arbor trees. Research results are as follows:
(1) There were differences in photosynthetic properties among different camellia species and different clones of C. oleifera. The order of daily average net photosynthetic efficiency of different camellia species was C. oleifera>C. osmantha-> C. gigantocarpa> C. vietnamensis> C. japonica> C. polyodonta. The daily average net photosynthetic efficiency of C.oleifera was the highest, at7.24μmol·m-2·s-1, and C.polyodonta was the lowest, at3.72μmol·m-2·s-1, merely taking up51.52%of C. oleifera. Among the tested C. oleifera clones, photosynthetic properties of Xianglin86was the highest, at6.86μmol·m-2·s-1, Cenruan2was the lowest at3.97μmol·m-2·s-1, only taking up57.87%of Xianglin86. The daily average net photosynthetic efficiency followed this order:Xianglin86> Cenruan24> Changlin4> Cenruan3> Gan190> Cenruan2, respectively.
(2) By simulating the light radiation condition, the maximum net photosynthetic efficiency of different camellia species ranged between8.4and10.5μmol·m-2·s-1, with C.oleifera the highest and C. polyodonta the lowest. Light compensation point of C.oleifera was merely15.50μmol·m-2·s-1, with the high capacity to utilize low light, followed by C. osmantha and C.japonica. C. polyodonta, C. vietnamensis and C. gigantocarpa had weak capacity to utilize low light. Light saturation point of C. osmantha which was distributed in areas of low latitude, showed high tolerance against strong light, while C.japonica which was distributed at high latitude showed the opposite. Among the tested clones, the maximum net photosynthetic efficiency ranged between5.8and11.4μmol·m-2·s-1, Cenruan3the highest and Cenruan2the lowest. The light compensation point of Gan190was16.36μmol·m-2·s-1,with the strongest capacity to utilize low light, while Cenruan2and Changlin4showed the weak capacity to utilize low light. Light saturation point of Cenruan3was537.86μmol·m-2·s-1, with the highest capacity to utilize strong light, while Gan190had the weakest capacity to utilize low light, with the light saturation point at383.55μmol·m-2·s-1.
(3) In late September and early October, under the shading-free condition, the curve of diurnal variation of net photosynthetic rate of the tested camellia species during seedling stage as well as C. oleifera clones during seedling and adult stage demonstrated single and double apexes. C. gigantocarpa of camellia species and Cenruan24of C. oleifera clone during seedling stage and Xianglin86during adult stage showed single apexes. The remaining camellia species, Sect. Paracamellia, C. vietnamensis and C. polyodonta during the seedling stage, and Cenruan2, Cenruan3, Xianglin86and Gan190of C. oleifera clones during the seedling stage, as well as Cenruan3and Gan190during the adult stage showed double apexes. Along with the increase of shading intensity, the curve of diurnal variation of net photosynthetic rate of all the camellia species during seedling stage and C. oleifera clones during seedling and adult stage showed single apexes. Under the condition of60%-90%shading, the curve of diurnal variation of net photosynthetic rate of all tested camellia species during seedling stage and C. oleifera clones during seedling and adult stage was lower than the control. Ratio of daily average net photosynthetic efficiency against that of control was higher than the ratio of photosynthetic active radiation against that of control under the same shading condition, indicating that under the condition of60%-90%shading, capacity of leaves of all tested camellia species and C.oleifera clones to capture low light was increased.
(4) Content of chlorophyll a, b and carotenoid in leaves of C. vietnamensis, C. oleifera, C. polyodonta, C. osmantha, Cenruan2, Cenruan3and Xianglin86during seedling stage and Cenruan3, Xianglin86and Gan190during adult stage under60%-90%shading condition increased obviously. Content of chlorophyll a, b and carotenoid in leaves of C. gigantocarpa, Cenruan24and Gan190under75%-90%shading condition was obviously higher than that of control. The synthesis of leaves chlorophyll and carotenoid under low light was enhanced in consistency with the increased capacity to utilize low light under shading condition for photosynthesis. Under low-light condition, increase of Qchlb/Qchla in the tested camellia species and C. oleifera clones during seedling stage was not obvious. During the process of chlorophyll synthesis, chlorophyll b was not synthesized in priority. Except C. oleifera, Qchlb/Qchla of remaining camellia species and C. oleifera was obviously higher than that of control, the increase extent of chlorophyll was higher than that of carotenoid, and chlorophyll was synthesized in priority. Among the C. oleifera clones during adult stage, except Qchlb/Qchla of Gan190under90%shading condition was obviously higher than that of control, Qchlb/Qchla of the remaining shading treatments was not increased significantly, and tendency to synthesize chlorophyll in priority was not obvious. However, Qchlb/Qchla in leaves of three tested clones under75%and90%shading treatments as well as Xianglin86and Gan190under60%shading treatment was increased significantly, chlorophyll b was synthesized in priority to better capture scattered light.
(5) During seedling stage, among the tested camellia species, the tolerance ability against low light of C. gigantocarpa and C. vietnamensis was the highest that there was no inhibiting to the growth of root, stem and leaves under90%shading treatment condition, and the growth of root was enhanced under the situation of60%shading intensity. C. oleifera and C. osmantha had relatively strong ability of low-light tolerance; there was no significant impact of each shading treatment on the growth of root, stem and leaves. C.polyodonta had relatively weak ability of low-light tolerance during seedling stage, and the growth of root system was inhibited obviously under60%shading treatment. Among the different clones of C. oleifera, Cenruan2and Gan190had the strongest ability of low-light tolerance, there was no inhibiting effects of each treatment on the growth of root, stem and leaves, and partial shading treatment enhanced the vegetative growth of the plant. Cenruan24had a relatively strong ability of low-light tolerance, although root growth was inhibited under90%shading condition, the growth of stem and leaves was enhanced under partial shading condition. Cenruan3and Xianglin86during seedling stage had a relatively weak ability of low-light tolerance, and75%shading would obviously inhibit the growth of root.
(6) Twenty-two months after shading treatment was applied on the tested C. oleifera clones during adult stage, it was found out that growth of Cenruan3and Xianglin86under90%shading condition and Gan190under60%-90%shading condition was prone to capturing light. Meanwhile,90%shading treatment also enhanced the growth of Xianglin86and Gan190towards capturing light. However, there was no significant difference in the content of N, P and K between three clones and control under each shading treatment. Under60%shading condition, Cenruan3had normal bud initiation and fruits, and there was no significant difference in fruit quality and oil composition compared with control. Though Gan190had similar bud quantity, fruit quantity and oil composition compared with control, fruit enlargement was inhibited. Bud initiation and fruits of Xianglin86were significantly lower than that of control. Although oil composition of fruits of Cenruan3, Xianglin86and Gan190under75%and90%shading condition was similar with that of control, bud initiation and fruits were obviously inhibited. At the adult stage, ability of low-light tolerance of three clones followed this order:Cenruan3>Gan190>Xianglin86.
(7) At seedling stage, camellia species and C. oleifera clones all had relatively strong ability of low-light tolerance. C. gigantocarpa, C. vietnamensis, C. oleifera, C. osmantha, Cenruan2and Gan190of C. oleifera could grow normally under90%shading condition (in early October photosynthetic active radiation was61.83μmol·m-2·s-1). Cenruan24could grow normally under75%shading condition (in early October photosynthetic active radiation was178.87μmol·m-2·s-1). Cenruan3and Xianglin86could also grow normally under60%shading condition (in early October photosynthetic active radiation was311.91μmol·m-2·s-1). C. oleifera clones at adult stage had relatively weak ability of low-light tolerance. Bud initiation and fruits of Cenruan3, Xianglin86and Gan190under75%shading condition (on September30th the diurnal average synthetic active radiation was193.25μmol·m-2·s-1) were inhibited. However, Cenruan3could grow normally under60%shading condition (on September30th the diurnal average synthetic active radiation was280.72μmol·m-2·s-1), and Gan190could also flower and have fruits normally.
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