冷凉地区生草制苹果园水分与光能利用特性及调控机制研究
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摘要
果树与环境之间的协调关系是实现果树产业可持续发展的重要途径,而土壤作为果树产业可持续发展的重要载体,其质量的维持至关重要。传统的土壤管理方式——清耕制严重破坏了表层土壤的生物进程,土壤有机质含量持续下降,降低了微生物群落结构的稳定性,从而导致土壤养分失衡、环境恶化,已成为制约果树产业可持续发展的主要瓶颈之一。果园生草在改善土壤环境和维持果园土壤肥力等方面具有重要的作用,已成为果品安全、优质持续生产的关键技术之一。光合作用是果树产量形成的基础,水分和光照既是影响光合作用的重要环境因子,也是树草共存系统相互影响最敏感的环境因子。因此,本研究以冷凉地区生草制苹果园为试材,研究了生草覆盖果园苹果植株水分与光能利用特性,以及遮阴和修剪(或刈割)条件下树草相互作用的生理生态机制,探讨了生草覆盖影响树体水分和光能利用的主要因子。主要结果如下：
1.生草覆盖和清耕处理寒富苹果叶片ψo、RC/CSo、ETo/CSo、PICS和△I均随着叶片光合能力的增强而升高,幼龄叶RC/CSo、ETo/CSo、PIcs和△I显著低于成龄叶；幼龄叶PSII的功能已有很大程度的发育。生草覆盖处理叶片Fv/Fm、PIcS和△I在5月7日显著高于清耕处理,说明生草覆盖处理叶片PSII和PSI的功能的发育早于清耕,因此具有较高的光合效率。
2.与清耕处理相比,自然高温强光条件下生草覆盖处理叶片AQY和Pm显著升高、ABS/RC和DIo/RC显著降低、Fv/Fm和Ψo显著升高,引起Plabs和△I显著升高。因此,高温强光下生草覆盖处理通过调控能量分配协调了PSI/PSII的关系,提高了PSI和PSII的活性,缓解了高温强光对整个光合机构的光抑制。
3.与清耕相比,生草覆盖处理寒富苹果叶片衰老过程中SOD和POD的活性平均高12.7%和11.4%,02·-和H202含量平均下降了21.2%和2.8%,减轻或推迟了膜脂过氧化的发生,减轻了叶绿体功能和结构的损伤。生草覆盖处理Fv/Fo和Fm在衰老中后期和后期均显著高于清耕,P。在中前期、中期和中后期均显著高于清耕。说明生草覆盖在一定程度上减轻或延迟了衰老对PSII结构和功能的损伤,提高了叶片制造的能力。
4.在冷凉地区的整个生长季节,行间生草区10cm土壤含水量(SWC)在6月、7月、9月和10月显著高于清耕,20cm SWC在6月～9月显著高于清耕,30cm SWC仅在6月显著高于清耕,而40cm SWC在8月～10月显著高于清耕,推测草域根系主要分布于0-30cm土层。株间覆盖区10cm SWC在6月及60cm SWC在6月～10月显著高于清耕,20cm SWC在5月～9月、30cm SWC在5月～8月及40cm SWC在7月和9月显著低于清耕,推测生草覆盖区苹果根系主要分布于0-40cm土层,清耕区苹果根系的主要分布层深于生草覆盖。说明果树根系与草根系水分相互作用的主要区域为0-40cm土层。
5.红三叶留茬20cm刈割时,茎叶光合效率较高、有效光合时间较长,组织纤维化程度和分蘖数居中、分枝数高、总产量高,有利于草域的持续生产、总产量的提高。因此,20cm为红三叶草的适宜刈割高度。
6.夏季晴天10：00气温迅速升高时,与覆草处理相比,‘覆草+遮阴’处理土壤温度低、升温慢,苹果叶片PSⅡ受体侧电子传递明显受阻,最大光化学效率和用于电子传递的量子效率显著降低；清耕处理土壤温度高、升温快,苹果叶片PSII受体侧电子传递明显受阻,用于电子传递的量子效率显著降低。覆草、‘覆草+遮阴’和清耕处理叶片净光合速率依次降低,覆草与清耕处理差异显著。说明生草制果园的覆盖厚度不宜过厚。‘覆草+遮阴’和覆草处理延迟了干旱和水分饱和胁迫条件下叶片光合效率的降低,促进了干旱复水后叶片光合效率的恢复。
7.干旱第4d,红三叶、早熟禾和清耕处理苹果叶片P。分别为干旱第1d的29.0%、9.3%和1.4%,Plabs分别为干旱第1d的68.6%、60.3%和28.5%,说明红三叶和早熟禾处理提升了寒富苹果对干旱胁迫的耐受性,红三叶处理的效果优于早熟禾。复水后第3d红三叶、早熟禾和清耕处理叶片P。恢复到干旱第1d的81.3%、80.1%和64.1%,说明红三叶和早熟禾处理有利于干旱复水后苹果叶片光合效率的恢复,红三叶的效果优于早熟禾。白三叶处理缓解了淹水对苹果叶片PSII放氧复合体和受体侧的伤害,提高了Fv/Fm和用于电子传递的量子效率,维持了PSII较高的性能,使叶片具有较高的光合效率；早熟禾处理的效果与白三叶处理相反。
Coordinating the relationship between the fruit trees and the environment is the important pathway to the sustainable development of fruit industry. Therefore, keeping the high quality of soil, as the important medium for the sustainable development of fruit industry, is essential. Bare earth, the traditional soil management system, severely broke the biological processes in topsoil. It also decreased the content of soil organic matter and stability of the microbial community structure, and then resulting in soil nutrient imbalances and environmental degradation which has become one of the major bottlenecks to the sustainable development of fruit industry. Considering the important function of maintaining soil fertility and improving soil environment, orchard grassing has become one of the key technologies of safe and high-quality continuous fruit production. Photosynthesis is the basis of the fruit yield. Moisture and light are not only the important environmental factors affecting photosynthesis, but also the most sensitive environmental factors of the coexistent system between fruit and grass.'Hanfu' apple orchard, with grass coverage in cool region, was used to study the effect of grass coverage on moisture and light utilization characteristics of'Hanfu' apple, and physiological and ecological mechanism of interactions between trees and grass under shading and pruning (or mowing) conditions. Major factors affecting water and solar energy utilization of 'Hanfu' apple tree under grassing condition are discussed. The main results are as follows:
1.ψo,φEo, RC/CSo, ETo/CSo, PIcs and ΔI were elevated with the enhanced photosynthetic capacity under grass coverage and bare earth conditions. RC/CSo, ETo/CSo, PICS and ΔI in young leaf were significantly higher than that in mature leaf under the two conditions. The function of PSII in young leaf have developed to a great extent. In May7, Fv/Fm, PIcs and ΔI under grass coverage condition were significant higher than that under bare earth. Development of PSII and PSI function under grass coverage were earlier than that under bare earth and therefore had a higher photosynthetic efficiency.
2. Compared to bare earth condition, AQY and Pm of 'Hanfu' apple leaf in natural high-temperature and high-light were significantly higher under grass coverage condition. The energy flux (per reaction centre) for absorption (ABS/RC) and dissipation (DIo/RC) significantly decreased. The maximal quantum yield of primary photochemistry (Fv/Fm) and and the efficiency (ψo) significantly increased which lead to the significant increase of performance index (PIabs). Therefore, grass coverage is able to coordinate the relationship between the PSI and PSII, improved the activity of PSI and PSII and relieved the photoinhibition to photosynthetic apparatus in high-temperature and high-light.
3. Compared to bare earth condition, the activity of SOD and POD in'Hanfu' apple leaf increased by an average of12.7%and11.4%, respectively, during leaf ageing under grass coverage. At the same time, the content of O2-and H2O2decreased by an average of21.2%and2.8%, respectively, which reduced or delayed the occurrence of membrane lipid peroxidation and mitigated the damage to function and structure of chloroplast. Fv/F0and Fm under grass coverage were significantly higher than that under bare earth at mid-late and late stage. Pn under grass coverage condition was significantly higher than that under bare earth at early-middle, middle and late-middle stage of leaf ageing. The results indicated that grass coverage, to a certain extent, could reduce or delay the damage of senescence on structure and function of PSⅡ. It also improved the leaf ability of manufacturing the assimilates,
4. From May to October, the soil water content (SWC) of10cm layer in alley grassing area in June, July, September and October were significantly higher than that in alley bare earth area, and SWC of20cm layer were significantly higher than that in alley bare earth from June to September. However, SWC of30cm layer were significantly higher than that in alley bare earth only in June, and SWC of40cm layer were significantly higher from August to October. It can be speculated that0-30cm topsoil was the main root distribution area of grass. SWC of10cm layer in June, and60cm layer from June to October in grass mulch area were significantly higher than that in bare earth area inline, while SWC of20cm layer from May to September,30cm alyer from May to August, and40cm layer in July and September significantly lower than that in bare earth area inliine. It can be inferred that0-40cm topsoil was the main apple root distribution area in grass mulch area, and the distribution of apple root were deeper in bare earth area than that in grass mulch area. The results suggested that the0-40cm topsoil was the main area, where existed the interaction for water between apple trees and grass.
5. Mowing for20cm of red clover stubble increased photosynthetic efficiency, prolong the effective photosynthetic time and increased branching number. In addition, the degree of fibrosis and the number of tillers of red clover were in the middle compared with the other treatments. Mowing for20cm stubble were beneficial to establishing the continuous production of grass group and improvement of total output. Therefore, mowing for20cm stubble was the suitable height for the planted red clover group.
6. Compared to the 'grass mulch'(GM) treatment, Soil temperature in'grass mulch+shade'(GM+S) treatment was too low and increased slowly, while the air temperature rose rapidly at10:00in summer.'GM+S'also accelerated the accumulation of QA-in acceptor side of PSⅡ and the significantly reduction of maximum quantum yield of primary photochemistry and quantum yield for electron transport. Soil temperature in 'bare earth' treatment (CK) was the highest among the three treatments and increased rapidly. The quantum yield for electron transport of apple leaf significantly decreased than that in GM due to the accumulation of QA-in acceptor side of PSII. The Pn of apple leaf were descending order in GM,'GM+S'and CK treatments, and the Pn was significant difference between GM and CK treatments. Therefore, the thickness of grass mulch shouldn't be too thick in grass coverage orchard. However,'GM+S'and'GM' treatments delay the period of the photosynthetic efficiency decline under drought and water saturation stress conditions, and promoted the recovery of the photosynthetic efficiency under rewatering after drought stress.
7. The Pn of apple leaf, at the4d after drought, were decreased into the29.0%,9.3%and1.4%of the Pn at the1d after drought in red clover, bluegrass and bare earth treatments, respectively, and PIabs were decreased into the68.6%,60.3%and28.5%of the PIabs at the1d after drought, respectively. This indicated the adaptability of apple trees to drought stress were improved by red clover and bluegrass treatments compared with the bare earth, while the adaptability in red clover treatment was better than that in bluegrass. At the3d after rewatering, Pn of apple leaf were recovered into the81.3%、80.1%and64.1%of the Pn at the1d after drought in red clover, bluegrass and clean tillage treatments, respectively. The results indicated that red clover and bluegrass treatments availed to the recovery of apple leaf photosynthetic efficiency in rewatering period, and the effect of red clover was better than that of bluegrass. White clover treatment alleviated the damage to oxygen evolving complex and the acceptor side of PSII, increased the maximum quantum yield of primary photochemistry and quantum yield for electron transport of PSII and maintained the higher performance of PSII during the flooding process. Finally, white clover treatment maintained the higher photosynthetic efficiency of apple leaf. The effect of bluegrass was opposite to the white clover.
1.生草覆盖和清耕处理寒富苹果叶片ψo、RC/CSo、ETo/CSo、PICS和△I均随着叶片光合能力的增强而升高,幼龄叶RC/CSo、ETo/CSo、PIcs和△I显著低于成龄叶；幼龄叶PSII的功能已有很大程度的发育。生草覆盖处理叶片Fv/Fm、PIcS和△I在5月7日显著高于清耕处理,说明生草覆盖处理叶片PSII和PSI的功能的发育早于清耕,因此具有较高的光合效率。
2.与清耕处理相比,自然高温强光条件下生草覆盖处理叶片AQY和Pm显著升高、ABS/RC和DIo/RC显著降低、Fv/Fm和Ψo显著升高,引起Plabs和△I显著升高。因此,高温强光下生草覆盖处理通过调控能量分配协调了PSI/PSII的关系,提高了PSI和PSII的活性,缓解了高温强光对整个光合机构的光抑制。
3.与清耕相比,生草覆盖处理寒富苹果叶片衰老过程中SOD和POD的活性平均高12.7%和11.4%,02·-和H202含量平均下降了21.2%和2.8%,减轻或推迟了膜脂过氧化的发生,减轻了叶绿体功能和结构的损伤。生草覆盖处理Fv/Fo和Fm在衰老中后期和后期均显著高于清耕,P。在中前期、中期和中后期均显著高于清耕。说明生草覆盖在一定程度上减轻或延迟了衰老对PSII结构和功能的损伤,提高了叶片制造的能力。
4.在冷凉地区的整个生长季节,行间生草区10cm土壤含水量(SWC)在6月、7月、9月和10月显著高于清耕,20cm SWC在6月～9月显著高于清耕,30cm SWC仅在6月显著高于清耕,而40cm SWC在8月～10月显著高于清耕,推测草域根系主要分布于0-30cm土层。株间覆盖区10cm SWC在6月及60cm SWC在6月～10月显著高于清耕,20cm SWC在5月～9月、30cm SWC在5月～8月及40cm SWC在7月和9月显著低于清耕,推测生草覆盖区苹果根系主要分布于0-40cm土层,清耕区苹果根系的主要分布层深于生草覆盖。说明果树根系与草根系水分相互作用的主要区域为0-40cm土层。
5.红三叶留茬20cm刈割时,茎叶光合效率较高、有效光合时间较长,组织纤维化程度和分蘖数居中、分枝数高、总产量高,有利于草域的持续生产、总产量的提高。因此,20cm为红三叶草的适宜刈割高度。
6.夏季晴天10：00气温迅速升高时,与覆草处理相比,‘覆草+遮阴’处理土壤温度低、升温慢,苹果叶片PSⅡ受体侧电子传递明显受阻,最大光化学效率和用于电子传递的量子效率显著降低；清耕处理土壤温度高、升温快,苹果叶片PSII受体侧电子传递明显受阻,用于电子传递的量子效率显著降低。覆草、‘覆草+遮阴’和清耕处理叶片净光合速率依次降低,覆草与清耕处理差异显著。说明生草制果园的覆盖厚度不宜过厚。‘覆草+遮阴’和覆草处理延迟了干旱和水分饱和胁迫条件下叶片光合效率的降低,促进了干旱复水后叶片光合效率的恢复。
7.干旱第4d,红三叶、早熟禾和清耕处理苹果叶片P。分别为干旱第1d的29.0%、9.3%和1.4%,Plabs分别为干旱第1d的68.6%、60.3%和28.5%,说明红三叶和早熟禾处理提升了寒富苹果对干旱胁迫的耐受性,红三叶处理的效果优于早熟禾。复水后第3d红三叶、早熟禾和清耕处理叶片P。恢复到干旱第1d的81.3%、80.1%和64.1%,说明红三叶和早熟禾处理有利于干旱复水后苹果叶片光合效率的恢复,红三叶的效果优于早熟禾。白三叶处理缓解了淹水对苹果叶片PSII放氧复合体和受体侧的伤害,提高了Fv/Fm和用于电子传递的量子效率,维持了PSII较高的性能,使叶片具有较高的光合效率；早熟禾处理的效果与白三叶处理相反。
Coordinating the relationship between the fruit trees and the environment is the important pathway to the sustainable development of fruit industry. Therefore, keeping the high quality of soil, as the important medium for the sustainable development of fruit industry, is essential. Bare earth, the traditional soil management system, severely broke the biological processes in topsoil. It also decreased the content of soil organic matter and stability of the microbial community structure, and then resulting in soil nutrient imbalances and environmental degradation which has become one of the major bottlenecks to the sustainable development of fruit industry. Considering the important function of maintaining soil fertility and improving soil environment, orchard grassing has become one of the key technologies of safe and high-quality continuous fruit production. Photosynthesis is the basis of the fruit yield. Moisture and light are not only the important environmental factors affecting photosynthesis, but also the most sensitive environmental factors of the coexistent system between fruit and grass.'Hanfu' apple orchard, with grass coverage in cool region, was used to study the effect of grass coverage on moisture and light utilization characteristics of'Hanfu' apple, and physiological and ecological mechanism of interactions between trees and grass under shading and pruning (or mowing) conditions. Major factors affecting water and solar energy utilization of 'Hanfu' apple tree under grassing condition are discussed. The main results are as follows:
1.ψo,φEo, RC/CSo, ETo/CSo, PIcs and ΔI were elevated with the enhanced photosynthetic capacity under grass coverage and bare earth conditions. RC/CSo, ETo/CSo, PICS and ΔI in young leaf were significantly higher than that in mature leaf under the two conditions. The function of PSII in young leaf have developed to a great extent. In May7, Fv/Fm, PIcs and ΔI under grass coverage condition were significant higher than that under bare earth. Development of PSII and PSI function under grass coverage were earlier than that under bare earth and therefore had a higher photosynthetic efficiency.
2. Compared to bare earth condition, AQY and Pm of 'Hanfu' apple leaf in natural high-temperature and high-light were significantly higher under grass coverage condition. The energy flux (per reaction centre) for absorption (ABS/RC) and dissipation (DIo/RC) significantly decreased. The maximal quantum yield of primary photochemistry (Fv/Fm) and and the efficiency (ψo) significantly increased which lead to the significant increase of performance index (PIabs). Therefore, grass coverage is able to coordinate the relationship between the PSI and PSII, improved the activity of PSI and PSII and relieved the photoinhibition to photosynthetic apparatus in high-temperature and high-light.
3. Compared to bare earth condition, the activity of SOD and POD in'Hanfu' apple leaf increased by an average of12.7%and11.4%, respectively, during leaf ageing under grass coverage. At the same time, the content of O2-and H2O2decreased by an average of21.2%and2.8%, respectively, which reduced or delayed the occurrence of membrane lipid peroxidation and mitigated the damage to function and structure of chloroplast. Fv/F0and Fm under grass coverage were significantly higher than that under bare earth at mid-late and late stage. Pn under grass coverage condition was significantly higher than that under bare earth at early-middle, middle and late-middle stage of leaf ageing. The results indicated that grass coverage, to a certain extent, could reduce or delay the damage of senescence on structure and function of PSⅡ. It also improved the leaf ability of manufacturing the assimilates,
4. From May to October, the soil water content (SWC) of10cm layer in alley grassing area in June, July, September and October were significantly higher than that in alley bare earth area, and SWC of20cm layer were significantly higher than that in alley bare earth from June to September. However, SWC of30cm layer were significantly higher than that in alley bare earth only in June, and SWC of40cm layer were significantly higher from August to October. It can be speculated that0-30cm topsoil was the main root distribution area of grass. SWC of10cm layer in June, and60cm layer from June to October in grass mulch area were significantly higher than that in bare earth area inline, while SWC of20cm layer from May to September,30cm alyer from May to August, and40cm layer in July and September significantly lower than that in bare earth area inliine. It can be inferred that0-40cm topsoil was the main apple root distribution area in grass mulch area, and the distribution of apple root were deeper in bare earth area than that in grass mulch area. The results suggested that the0-40cm topsoil was the main area, where existed the interaction for water between apple trees and grass.
5. Mowing for20cm of red clover stubble increased photosynthetic efficiency, prolong the effective photosynthetic time and increased branching number. In addition, the degree of fibrosis and the number of tillers of red clover were in the middle compared with the other treatments. Mowing for20cm stubble were beneficial to establishing the continuous production of grass group and improvement of total output. Therefore, mowing for20cm stubble was the suitable height for the planted red clover group.
6. Compared to the 'grass mulch'(GM) treatment, Soil temperature in'grass mulch+shade'(GM+S) treatment was too low and increased slowly, while the air temperature rose rapidly at10:00in summer.'GM+S'also accelerated the accumulation of QA-in acceptor side of PSⅡ and the significantly reduction of maximum quantum yield of primary photochemistry and quantum yield for electron transport. Soil temperature in 'bare earth' treatment (CK) was the highest among the three treatments and increased rapidly. The quantum yield for electron transport of apple leaf significantly decreased than that in GM due to the accumulation of QA-in acceptor side of PSII. The Pn of apple leaf were descending order in GM,'GM+S'and CK treatments, and the Pn was significant difference between GM and CK treatments. Therefore, the thickness of grass mulch shouldn't be too thick in grass coverage orchard. However,'GM+S'and'GM' treatments delay the period of the photosynthetic efficiency decline under drought and water saturation stress conditions, and promoted the recovery of the photosynthetic efficiency under rewatering after drought stress.
7. The Pn of apple leaf, at the4d after drought, were decreased into the29.0%,9.3%and1.4%of the Pn at the1d after drought in red clover, bluegrass and bare earth treatments, respectively, and PIabs were decreased into the68.6%,60.3%and28.5%of the PIabs at the1d after drought, respectively. This indicated the adaptability of apple trees to drought stress were improved by red clover and bluegrass treatments compared with the bare earth, while the adaptability in red clover treatment was better than that in bluegrass. At the3d after rewatering, Pn of apple leaf were recovered into the81.3%、80.1%and64.1%of the Pn at the1d after drought in red clover, bluegrass and clean tillage treatments, respectively. The results indicated that red clover and bluegrass treatments availed to the recovery of apple leaf photosynthetic efficiency in rewatering period, and the effect of red clover was better than that of bluegrass. White clover treatment alleviated the damage to oxygen evolving complex and the acceptor side of PSII, increased the maximum quantum yield of primary photochemistry and quantum yield for electron transport of PSII and maintained the higher performance of PSII during the flooding process. Finally, white clover treatment maintained the higher photosynthetic efficiency of apple leaf. The effect of bluegrass was opposite to the white clover.
引文
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