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Functional Plant Biology Functional Plant Biology Society
Plant function and evolutionary biology
RESEARCH ARTICLE

Effects of contrasting shade treatments on the carbon production and antioxidant activities of soybean plants

Muhammad Ali Raza A , Ling Yang Feng A , Nasir Iqbal A , Imran Khan B , Tehseen Ahmad Meraj A , Zeng Jin Xi A , Muhammd Naeem A , Saeed Ahmed C , Muhammad Tayyab Sattar A D , Yuan Kai Chen A , Chen Hui Huan A , Mukhtar Ahmed E F , Feng Yang A G and Wenyu Yang A G
+ Author Affiliations
- Author Affiliations

A College of Agronomy, Sichuan Agricultural University, Chengdu 611130, PR China.

B Department of Grassland Science, Sichuan Agricultural University, Chengdu, 611130, PR China.

C College of Food Science, Sichuan Agricultural University, Yaan 625014, PR China.

D Institute of Ecological and Environmental Sciences, Sichuan Agricultural University, Chengdu, 611130, PR China.

E Department of Agronomy, Pir Mehr Ali Shah Arid Agriculture University, Rawalpindi, Pakistan.

F Department of Agricultural Research for Northern Sweden, Swedish University of Agricultural Sciences Umea, Sweden.

G Correspondending authors. Email: f.yang@sicau.edu.cn; mssiyangwy@sicau.edu.cn

Functional Plant Biology 47(4) 342-354 https://doi.org/10.1071/FP19213
Submitted: 2 August 2019  Accepted: 24 November 2019   Published: 11 February 2020

Abstract

In China, maize-soybean relay-intercropping system follow the two main planting-patterns: (i) traditional relay-intercropping; maize-soybean equal row planting, where soybean experience severe maize shading on both sides of plants, and (ii) modern relay-intercropping; narrow-wide row planting, in this new planting pattern only one side of soybean leaves suffer from maize shading. Therefore, in this study, changes in morphological traits, cytochrome content, photosynthetic characteristics, carbon status, and the activities of superoxide dismutase (SOD), peroxidase (POD), catalase (CAT) and ascorbate peroxidase (APX) were investigated at 30 days after treatment (DAT) in shade-tolerant soybean variety Nandou-12 subjected to three different types of shading conditions; normal light (NL, all trifoliate-leaves of soybean plants were under normal light); unilateral shade (US, all right-side trifoliate-leaves of soybean plants from top to bottom were under shade while all the left-side of trifoliate-leaves from top to bottom were in normal light); bilateral shade (BS, all trifoliate-leaves of soybean plants were under complete shade). Compared with BS, US conditions decreased plant height and increased stem diameter, leaf area, and biomass at 30 DAT. Biomass distribution rates to stem, petiole and leaves, and photosynthetic characteristics were markedly improved by the US at all sampling stages, which proved to be a better growing condition than BS with respect to shade tolerance. The enhanced net photosynthesis and transpiration rates in the left-side leaves (LS) of soybean plants in US, when compared with the LS in BS, allowed them to produce higher total soluble sugar (by 70%) and total soluble protein (by 17%) at 30 DAT which reduce the adverse effects of shading at right-side leaves (RS) of the soybean plants. Similarly, soybean leaves under US accumulated higher proline content in US than the leaves of BS plants. Soybean leaves grown in shading conditions (LS and RS of BS and RS of US) developed antioxidative defence-mechanisms, including the accelerated activities of SOD, POD, APX, and CAT. Comparatively, soybean leaves in US displayed lower activity levels of the antioxidative enzymes than the leaves of BS plants, showing that soybean plants experienced less shade stress in US as compared with BS treatment. Overall, these results indicate that the association of improved photosynthetic characteristics, sugar and protein accumulation and optimum antioxidative defences could be an effective approach for growing soybean in intercropping environments.

Additional keywords: antioxidants, chlorophyll, photosynthesis, protein content.


References

AbdElgawad H, Farfan-Vignolo ER, De Vos D, Asard H (2015) Elevated CO2 mitigates drought and temperature-induced oxidative stress differently in grasses and legumes. Plant Science 231, 1–10.
Elevated CO2 mitigates drought and temperature-induced oxidative stress differently in grasses and legumes.Crossref | GoogleScholarGoogle Scholar | 25575986PubMed |

Ahmed S, Raza M, Zhou T, Hussain S, Khalid M, Feng L, Wasaya A, Iqbal N, Ahmed A, Liu W (2018) Responses of soybean dry matter production, phosphorus accumulation, and seed yield to sowing time under relay intercropping with maize. Agronomy 8, 282
Responses of soybean dry matter production, phosphorus accumulation, and seed yield to sowing time under relay intercropping with maize.Crossref | GoogleScholarGoogle Scholar |

Amiard V, Mueh KE, Demmig-Adams B, Ebbert V, Turgeon R, Adams WW (2005) Anatomical and photosynthetic acclimation to the light environment in species with differing mechanisms of phloem loading. Proceedings of the National Academy of Sciences of the United States of America 102, 12968–12973.
Anatomical and photosynthetic acclimation to the light environment in species with differing mechanisms of phloem loading.Crossref | GoogleScholarGoogle Scholar | 16120679PubMed |

Amirjani M (2010) Effect of salinity stress on growth, mineral composition, proline content, antioxidant enzymes of soybean. American Journal of Plant Physiology 5, 350–360.
Effect of salinity stress on growth, mineral composition, proline content, antioxidant enzymes of soybean.Crossref | GoogleScholarGoogle Scholar |

Apel K, Hirt H (2004) Reactive oxygen species: metabolism, oxidative stress, and signal transduction. Annual Review of Plant Biology 55, 373–399.
Reactive oxygen species: metabolism, oxidative stress, and signal transduction.Crossref | GoogleScholarGoogle Scholar | 15377225PubMed |

Asada K (1999) The water-water cycle in chloroplasts: scavenging of active oxygens and dissipation of excess photons. Annual Review of Plant Biology 50, 601–639.
The water-water cycle in chloroplasts: scavenging of active oxygens and dissipation of excess photons.Crossref | GoogleScholarGoogle Scholar |

Ballaré CL, Pierik R (2017) The shade‐avoidance syndrome: multiple signals and ecological consequences. Plant, Cell & Environment 40, 2530–2543.
The shade‐avoidance syndrome: multiple signals and ecological consequences.Crossref | GoogleScholarGoogle Scholar |

Bates L, Waldren R, Teare I (1973) Rapid determination of free proline for water-stress studies. Plant and Soil 39, 205–207.
Rapid determination of free proline for water-stress studies.Crossref | GoogleScholarGoogle Scholar |

Blokhina O, Virolainen E, Fagerstedt KV (2003) Antioxidants, oxidative damage and oxygen deprivation stress: a review. Annals of Botany 91, 179–194.
Antioxidants, oxidative damage and oxygen deprivation stress: a review.Crossref | GoogleScholarGoogle Scholar | 12509339PubMed |

Bradford MM (1976) A rapid and sensitive method for the quantitation of microgram quantities of protein utilizing the principle of protein-dye binding. Analytical Biochemistry 72, 248–254.
A rapid and sensitive method for the quantitation of microgram quantities of protein utilizing the principle of protein-dye binding.Crossref | GoogleScholarGoogle Scholar | 942051PubMed |

Chaki M, Fernández-Ocaña AM, Valderrama R, Carreras A, Esteban FJ, Luque F, Gómez-Rodríguez MV, Begara-Morales JC, Corpas FJ, Barroso JB (2008) Involvement of reactive nitrogen and oxygen species (RNS and ROS) in sunflower–mildew interaction. Plant & Cell Physiology 50, 265–279.
Involvement of reactive nitrogen and oxygen species (RNS and ROS) in sunflower–mildew interaction.Crossref | GoogleScholarGoogle Scholar |

Corpas FJ, Chaki M, Fernandez-Ocana A, Valderrama R, Palma JM, Carreras A, Begara-Morales JC, Airaki M, del Río LA, Barroso JB (2008) Metabolism of reactive nitrogen species in pea plants under abiotic stress conditions. Plant & Cell Physiology 49, 1711–1722.
Metabolism of reactive nitrogen species in pea plants under abiotic stress conditions.Crossref | GoogleScholarGoogle Scholar |

Echer FR, Rosolem CA (2015) Cotton leaf gas exchange responses to irradiance and leaf aging. Biologia Plantarum 59, 366–372.
Cotton leaf gas exchange responses to irradiance and leaf aging.Crossref | GoogleScholarGoogle Scholar |

Fan Y, Chen J, Cheng Y, Raza MA, Wu X, Wang Z, Liu Q, Wang R, Wang X, Yong T (2018) Effect of shading and light recovery on the growth, leaf structure, and photosynthetic performance of soybean in a maize-soybean relay-strip intercropping system. PLoS One 13, e0198159
Effect of shading and light recovery on the growth, leaf structure, and photosynthetic performance of soybean in a maize-soybean relay-strip intercropping system.Crossref | GoogleScholarGoogle Scholar | 30379963PubMed |

Fehr WR (1977) Stages of soybean development. Kenya soil survey. Executive report no. 80. Iowa State University Co-operative and Extension Service, Ames, IA, USA.

Feng LY, Raza MA, Chen Y, Khalid MHB, Meraj TA, Ahsan F, Fan Y, Du J, Wu X, Song C (2019) Narrow-wide row planting pattern improves the light environment and seed yields of intercrop species in relay intercropping system. PLoS One 14, e0212885

Feng LY, Raza MA, Li ZC, Chen Y, Khalid MHB, Du J, Liu W, Wu X, Song C, Yu L (2019) The influence of light intensity and leaf movement on photosynthesis characteristics and carbon balance of soybean. Frontiers in Plant Science 9, 1952
The influence of light intensity and leaf movement on photosynthesis characteristics and carbon balance of soybean.Crossref | GoogleScholarGoogle Scholar |

Gommers CM, Visser EJ, St Onge KR, Voesenek LA, Pierik R (2013) Shade tolerance: when growing tall is not an option. Trends in Plant Science 18, 65–71.
Shade tolerance: when growing tall is not an option.Crossref | GoogleScholarGoogle Scholar | 23084466PubMed |

Grace SC, Logan BA (1996) Acclimation of foliar antioxidant systems to growth irradiance in three broad-leaved evergreen species. Plant Physiology 112, 1631–1640.
Acclimation of foliar antioxidant systems to growth irradiance in three broad-leaved evergreen species.Crossref | GoogleScholarGoogle Scholar | 12226469PubMed |

Hansen J, Møller I (1975) Percolation of starch and soluble carbohydrates from plant tissue for quantitative determination with anthrone. Analytical Biochemistry 68, 87–94.
Percolation of starch and soluble carbohydrates from plant tissue for quantitative determination with anthrone.Crossref | GoogleScholarGoogle Scholar | 1190454PubMed |

Hoagland, DR, Arnon, DI (1938) ‘Growing plants without soil by the water-culture method.’ (University of California, College of Agriculture: Agricultural Experiment Station, Berkeley, CA, USA)

Iqbal N, Hussain S, Raza MA, Yang C, Safdar ME, Brestic M, Aziz A, Hayyat MS, Asghar MA, Yang W (2019) Drought tolerance of soybean (Glycine max L. Merr.) by improved photosynthetic characteristics and an efficient antioxidant enzyme system under a split-root system. Frontiers in Physiology 10, 786
Drought tolerance of soybean (Glycine max L. Merr.) by improved photosynthetic characteristics and an efficient antioxidant enzyme system under a split-root system.Crossref | GoogleScholarGoogle Scholar | 31333479PubMed |

Khalid M, Raza M, Yu H, Sun F, Zhang Y, Lu F, Si L, Iqbal N, Khan I, Fu F (2019) Effect of shade treatments on morphology, photosynthetic and chlorophyll fluorescence characteristics of soybeans (Glycine max L. Merr.). Applied Ecology and Environmental Research 17, 2551–2569.
Effect of shade treatments on morphology, photosynthetic and chlorophyll fluorescence characteristics of soybeans (Glycine max L. Merr.).Crossref | GoogleScholarGoogle Scholar |

Kurepin LV, Park JM, Lazarovits G, Hüner NP (2015) Involvement of plant stress hormones in Burkholderia phytofirmans-induced shoot and root growth promotion. Plant Growth Regulation 77, 179–187.
Involvement of plant stress hormones in Burkholderia phytofirmans-induced shoot and root growth promotion.Crossref | GoogleScholarGoogle Scholar |

Lawlor DW, Tezara W (2009) Causes of decreased photosynthetic rate and metabolic capacity in water-deficient leaf cells: a critical evaluation of mechanisms and integration of processes. Annals of Botany 103, 561–579.
Causes of decreased photosynthetic rate and metabolic capacity in water-deficient leaf cells: a critical evaluation of mechanisms and integration of processes.Crossref | GoogleScholarGoogle Scholar | 19155221PubMed |

Li R, Wen T, Tang Y, Sun X, Xia C (2014a) Effect of shading on photosynthetic and chlorophyll fluorescence characteristics of soybean. Caoye Xuebao 23, 198–206.

Li T, Liu L-N, Jiang C-D, Liu Y-J, Shi L (2014b) Effects of mutual shading on the regulation of photosynthesis in field-grown sorghum. Journal of Photochemistry and Photobiology. B, Biology 137, 31–38.
Effects of mutual shading on the regulation of photosynthesis in field-grown sorghum.Crossref | GoogleScholarGoogle Scholar | 24935099PubMed |

Liao J-X, Ge Y, Huang C-C, Zhang J, Liu Q-X, Chang J (2005) Effects of irradiance on photosynthetic characteristics and growth of Mosla chinensis and M. scabra. Photosynthetica 43, 111–115.
Effects of irradiance on photosynthetic characteristics and growth of Mosla chinensis and M. scabra.Crossref | GoogleScholarGoogle Scholar |

Lichtenthaler, HK, Buschmann, C (2001) ‘Chlorophylls and carotenoids: measurement and characterization by UV‐VIS spectroscopy. Current protocols in food analytical chemistry.’ (John Wiley & Sons Inc.: Hoboken, NJ, USA)

Lichtenthaler H, Buschmann C, Döll M, Fietz H-J, Bach T, Kozel U, Meier D, Rahmsdorf U (1981) Photosynthetic activity, chloroplast ultrastructure, and leaf characteristics of high-light and low-light plants and of sun and shade leaves. Photosynthesis Research 2, 115–141.
Photosynthetic activity, chloroplast ultrastructure, and leaf characteristics of high-light and low-light plants and of sun and shade leaves.Crossref | GoogleScholarGoogle Scholar | 24470202PubMed |

Lichtenthaler HK, Ač A, Marek MV, Kalina J, Urban O (2007) Differences in pigment composition, photosynthetic rates and chlorophyll fluorescence images of sun and shade leaves of four tree species. Plant Physiology and Biochemistry 45, 577–588.
Differences in pigment composition, photosynthetic rates and chlorophyll fluorescence images of sun and shade leaves of four tree species.Crossref | GoogleScholarGoogle Scholar | 17587589PubMed |

Lin J-S, Wang G-X (2002) Doubled CO2 could improve the drought tolerance better in sensitive cultivars than in tolerant cultivars in spring wheat. Plant Science 163, 627–637.
Doubled CO2 could improve the drought tolerance better in sensitive cultivars than in tolerant cultivars in spring wheat.Crossref | GoogleScholarGoogle Scholar |

Logan B, Barker D, Demmig‐Adams B, Adams WW (1996) Acclimation of leaf carotenoid composition and ascorbate levels to gradients in the light environment within an Australian rainforest. Plant, Cell & Environment 19, 1083–1090.
Acclimation of leaf carotenoid composition and ascorbate levels to gradients in the light environment within an Australian rainforest.Crossref | GoogleScholarGoogle Scholar |

Logan BA, Demmig-Adam B, Adams WW (1998) Antioxidants and xanthophyll cycle-dependent energy dissipation in Cucurbita pepo L. and Vinca major L. upon a sudden increase in growth PPFD in the field. Journal of Experimental Botany 49, 1881–1888.
Antioxidants and xanthophyll cycle-dependent energy dissipation in Cucurbita pepo L. and Vinca major L. upon a sudden increase in growth PPFD in the field.Crossref | GoogleScholarGoogle Scholar |

McCarthy-Suárez I, Gómez M, Del Río L, Palma J (2011) Role of peroxisomes in the oxidative injury induced by 2,4-dichlorophenoxyacetic acid in leaves of pea plants. Biologia Plantarum 55, 485–492.
Role of peroxisomes in the oxidative injury induced by 2,4-dichlorophenoxyacetic acid in leaves of pea plants.Crossref | GoogleScholarGoogle Scholar |

Miller G, Suzuki N, Ciftci‐Yilmaz S, Mittler R (2010) Reactive oxygen species homeostasis and signalling during drought and salinity stresses. Plant, Cell & Environment 33, 453–467.
Reactive oxygen species homeostasis and signalling during drought and salinity stresses.Crossref | GoogleScholarGoogle Scholar |

Park Y, Runkle ES (2017) Far-red radiation promotes growth of seedlings by increasing leaf expansion and whole-plant net assimilation. Environmental and Experimental Botany 136, 41–49.
Far-red radiation promotes growth of seedlings by increasing leaf expansion and whole-plant net assimilation.Crossref | GoogleScholarGoogle Scholar |

Porra R, Thompson W, Kriedemann P (1989) Determination of accurate extinction coefficients and simultaneous equations for assaying chlorophylls a and b extracted with four different solvents: verification of the concentration of chlorophyll standards by atomic absorption spectroscopy. Biochimica et Biophysica Acta (BBA) - Bioenergetics 975, 384–394.
Determination of accurate extinction coefficients and simultaneous equations for assaying chlorophylls a and b extracted with four different solvents: verification of the concentration of chlorophyll standards by atomic absorption spectroscopy.Crossref | GoogleScholarGoogle Scholar |

Possart A, Fleck C, Hiltbrunner A (2014) Shedding (far-red) light on phytochrome mechanisms and responses in land plants. Plant Science 217–218, 36–46.
Shedding (far-red) light on phytochrome mechanisms and responses in land plants.Crossref | GoogleScholarGoogle Scholar | 24467894PubMed |

Raza M, Feng L, Iqbal N, Manaf A, Khalid M, Wasaya A, Ansar M, Billah M, Yang F, Yang W (2018) Effect of sulphur application on photosynthesis and biomass accumulation of sesame varieties under rainfed conditions. Agronomy 8, 149
Effect of sulphur application on photosynthesis and biomass accumulation of sesame varieties under rainfed conditions.Crossref | GoogleScholarGoogle Scholar |

Raza MA, Feng LY, Iqbal N, Ahmed M, Chen YK, Khalid MHB, Din AMU, Khan A, Ijaz W, Hussain A (2019a) Growth and development of soybean under changing light environments in relay intercropping system. PeerJ 7, e7262
Growth and development of soybean under changing light environments in relay intercropping system.Crossref | GoogleScholarGoogle Scholar | 31372317PubMed |

Raza MA, Feng LY, Khalid MH, Iqbal N, Meraj TA, Hassan MJ, Ahmed S, Chen YK, Feng Y, Wenyu Y (2019b) Optimum leaf excision increases the biomass accumulation and seed yield of maize plants under different planting patterns. Annals of Applied Biology 175, 54–68.
Optimum leaf excision increases the biomass accumulation and seed yield of maize plants under different planting patterns.Crossref | GoogleScholarGoogle Scholar |

Raza MA, Feng LY, van der Werf W, Cai GR, Khalid MHB, Iqbal N, Hassan MJ, Meraj TA, Naeem M, Khan I (2019c) Narrow‐wide‐row planting pattern increases the radiation use efficiency and seed yield of intercrop species in relay‐intercropping system. Food and Energy Security 8, e170
Narrow‐wide‐row planting pattern increases the radiation use efficiency and seed yield of intercrop species in relay‐intercropping system.Crossref | GoogleScholarGoogle Scholar |

Raza MA, Feng LY, van der Werf W, Iqbal N, Khalid MHB, Chen YK, Wasaya A, Ahmed S, Din AMU, Khan A (2019d) Maize leaf-removal: a new agronomic approach to increase dry matter, flower number and seed-yield of soybean in maize soybean relay intercropping system. Scientific Reports 9, 13453
Maize leaf-removal: a new agronomic approach to increase dry matter, flower number and seed-yield of soybean in maize soybean relay intercropping system.Crossref | GoogleScholarGoogle Scholar | 31530859PubMed |

Raza MA, Feng LY, van der Werf W, Iqbal N, Khan I, Hassan MJ, Ansar M, Chen YK, Xi ZJ, Shi JY (2019e) Optimum leaf defoliation: a new agronomic approach for increasing nutrient uptake and land equivalent ratio of maize soybean relay intercropping system. Field Crops Research 244, 107647
Optimum leaf defoliation: a new agronomic approach for increasing nutrient uptake and land equivalent ratio of maize soybean relay intercropping system.Crossref | GoogleScholarGoogle Scholar |

Raza MA, Khalid MHB, Zhang X, Feng LY, Khan I, Hassan MJ, Ahmed M, Ansar M, Chen YK, Fan YF (2019f) Effect of planting patterns on yield, nutrient accumulation and distribution in maize and soybean under relay intercropping systems. Scientific Reports 9, 4947
Effect of planting patterns on yield, nutrient accumulation and distribution in maize and soybean under relay intercropping systems.Crossref | GoogleScholarGoogle Scholar | 30894625PubMed |

Rhizopoulou S, Meletiou-Christou M, Diamantoglou S (1991) Water relations for sun and shade leaves of four Mediterranean evergreen sclerophylls. Journal of Experimental Botany 42, 627–635.
Water relations for sun and shade leaves of four Mediterranean evergreen sclerophylls.Crossref | GoogleScholarGoogle Scholar |

Riazi A, Matsuda K, Arslan A (1985) Water-stress induced changes in concentrations of proline and other solutes in growing regions of young barley leaves. Journal of Experimental Botany 36, 1716–1725.
Water-stress induced changes in concentrations of proline and other solutes in growing regions of young barley leaves.Crossref | GoogleScholarGoogle Scholar |

Seemann JR (1989) Light adaptation/acclimation of photosynthesis and the regulation of ribulose-1,5-bisphosphate carboxylase activity in sun and shade plants. Plant Physiology 91, 379–386.
Light adaptation/acclimation of photosynthesis and the regulation of ribulose-1,5-bisphosphate carboxylase activity in sun and shade plants.Crossref | GoogleScholarGoogle Scholar | 16667029PubMed |

Siringam K, Juntawong N, Cha-um S, Kirdmanee C (2011) Salt stress induced ion accumulation, ion homeostasis, membrane injury and sugar contents in salt-sensitive rice (Oryza sativa L. spp. indica) roots under isoosmotic conditions. African Journal of Biotechnology 10, 1340–1346.

Slewinski TL, Braun DM (2010) Current perspectives on the regulation of whole-plant carbohydrate partitioning. Plant Science 178, 341–349.
Current perspectives on the regulation of whole-plant carbohydrate partitioning.Crossref | GoogleScholarGoogle Scholar |

Smirnoff N (2005) ‘Antioxidants and reactive oxygen species in plants.’ (John Wiley & Sons Inc: Hoboken, NJ, USA)

Valderrama R, Corpas FJ, Carreras A, Fernández-Ocaña A, Chaki M, Luque F, Gómez-Rodríguez MV, Colmenero-Varea P, Luis A, Barroso JB (2007) Nitrosative stress in plants. Febs Letters 581, 453–461.

Valladares F, Niinemets Ü (2008) Shade tolerance, a key plant feature of complex nature and consequences. Annual Review of Ecology Evolution and Systematics 39, 237–257.
Shade tolerance, a key plant feature of complex nature and consequences.Crossref | GoogleScholarGoogle Scholar |

Verhoeven AS, Swanberg A, Thao M, Whiteman J (2005) Seasonal changes in leaf antioxidant systems and xanthophyll cycle characteristics in Taxus × media growing in sun and shade environments. Physiologia Plantarum 123, 428–434.
Seasonal changes in leaf antioxidant systems and xanthophyll cycle characteristics in Taxus × media growing in sun and shade environments.Crossref | GoogleScholarGoogle Scholar |

Wu Y, Gong W, Yang W (2017) Shade inhibits leaf size by controlling cell proliferation and enlargement in soybean. Scientific Reports 7, 9259
Shade inhibits leaf size by controlling cell proliferation and enlargement in soybean.Crossref | GoogleScholarGoogle Scholar | 28835715PubMed |

Xie Y, Liu Y, Wang H, Ma X, Wang B, Wu G, Wang H (2017) Phytochrome-interacting factors directly suppress MIR156 expression to enhance shade-avoidance syndrome in Arabidopsis. Nature Communications 8, 348
Phytochrome-interacting factors directly suppress MIR156 expression to enhance shade-avoidance syndrome in Arabidopsis.Crossref | GoogleScholarGoogle Scholar | 28839125PubMed |

Xu C, Yin Y, Cai R, Wang P, Ni Y, Guo J, Chen E, Cai T, Cui Z, Liu T (2013) Responses of photosynthetic characteristics and antioxidative metabolism in winter wheat to post-anthesis shading. Photosynthetica 51, 139–150.
Responses of photosynthetic characteristics and antioxidative metabolism in winter wheat to post-anthesis shading.Crossref | GoogleScholarGoogle Scholar |

Yan Y, Gong W, Yang W, Wan Y, Chen X, Chen Z, Wang L (2010) Seed treatment with uniconazole powder improves soybean seedling growth under shading by corn in relay strip intercropping system. Plant Production Science 13, 367–374.
Seed treatment with uniconazole powder improves soybean seedling growth under shading by corn in relay strip intercropping system.Crossref | GoogleScholarGoogle Scholar |

Yang F, Huang S, Gao R, Liu W, Yong T, Wang X, Wu X, Yang W (2014) Growth of soybean seedlings in relay strip intercropping systems in relation to light quantity and red : far-red ratio. Field Crops Research 155, 245–253.
Growth of soybean seedlings in relay strip intercropping systems in relation to light quantity and red : far-red ratio.Crossref | GoogleScholarGoogle Scholar |

Yang F, Feng L, Liu Q, Wu X, Fan Y, Raza MA, Cheng Y, Chen J, Wang X, Yong T (2018) Effect of interactions between light intensity and red-to-far-red ratio on the photosynthesis of soybean leaves under shade condition. Environmental and Experimental Botany 150, 79–87.
Effect of interactions between light intensity and red-to-far-red ratio on the photosynthesis of soybean leaves under shade condition.Crossref | GoogleScholarGoogle Scholar |

Zheng Y, Mai B, Wu R, Feng Y, Sofo A, Ni Y, Sun J, Li J, Xu J (2011) Acclimation of winter wheat (Triticum aestivum, cv. Yangmai 13) to low levels of solar irradiance. Photosynthetica 49, 426
Acclimation of winter wheat (Triticum aestivum, cv. Yangmai 13) to low levels of solar irradiance.Crossref | GoogleScholarGoogle Scholar |

Zhu H, Li X, Zhai W, Liu Y, Gao Q, Liu J, Ren L, Chen H, Zhu Y (2017) Effects of low light on photosynthetic properties, antioxidant enzyme activity, and anthocyanin accumulation in purple pak-choi (Brassica campestris ssp. Chinensis Makino). PLoS One 12, e0179305
Effects of low light on photosynthetic properties, antioxidant enzyme activity, and anthocyanin accumulation in purple pak-choi (Brassica campestris ssp. Chinensis Makino).Crossref | GoogleScholarGoogle Scholar | 29240840PubMed |