Intercropping induces physiological and morphological plasticity in oilseed rape and barley under drought stress


  • Noushin SADEGHZADEH Department of Plant, Cell and Molecular Biology, Faculty of Natural Sciences, University of Tabriz, Tabriz, Iran
  • Roghieh HAJIBOLAND Department of Plant, Cell and Molecular Biology, Faculty of Natural Sciences, University of Tabriz, Tabriz, Iran
  • Charlotte POSCHENRIEDER Plant Physiology Laboratory, Bioscience Faculty, Universidad Autónoma de Barcelona, Spain



drought stress, intercropping, osmotic adjustment, photosynthesis rate, , transpiration, water use efficiency


Intercropping is an agricultural practice that can improve crop yield due to better availability of resources, including water. There are few studies, however, addressing the physiological mechanisms behind this phenomenon. In this work oilseed rape (Brassica napus L.) and barley (Hordeum vulgare L.) were cultivated either as monocrop (MC) or intercrop (IC) under well-watered (WW) or drought stress (DS) conditions in a growth chamber. After eight weeks DS, the leaf relative water content was higher in the IC compared with the MC plants in both species and the DS-induced senescence of old leaves was considerably postponed in oilseed rape. Intercropped oilseed rape showed elevated levels of leaf photosynthesis rate, superior accumulation of organic osmolytes but higher water loss compared with the MC counterparts under DS conditions. In barley, less transpiration, an increased root : shoot ratio and osmolyte accumulation was observed in the IC compared with MC plants under DS conditions. The water use efficiency was higher in the IC compared to MC barley and the plants yield was higher in the IC than in the MC oilseed rape. Our data showed that intercropping is a reliable practice for cultivation of both species under arid and semi-arid regions or under rainfed conditions.


Brooker, R. W., Bennett, A. E., Cong, W. F., Daniell, T. J., George, T. S., Hallett, P. D., Hawes, C., Iannetta, P. P., Jones, H. G., Karley, A. J., & Li, L. (2015). Improving intercropping: a synthesis of research in agronomy, plant physiology and ecology. New Phytologist, 206(1), 107‒117.

Chastain, D.R., Snider, J.L., Choinski, J.S., Collins, G.D., Perry, C.D., Whitaker, J., Grey, T.L., Sorensen, R. B., van Iersel, M., Byrd, S.A., & Porter, W. (2016). Leaf ontogeny strongly influences photosynthetic tolerance to drought and high temperature in Gossypium hirsutum. Journal of Plant Physiology, 199, 18‒28.

Chaves, M. M., Flexas, J., & Pinheiro, C. (2009). Photosynthesis under drought and salt stress: regulation mechanisms from whole plant to cell. Annals of Botany, 103, 551–560.

Chen, B. J., Hajiboland, R., Bahrami-Rad, S., Moradtalab, N., & Anten, N. P. (2019). Presence of belowground neighbors activates defense pathways at the expense of growth in tobacco plants. Frontiers in Plant Science, 10, 751.

Cruz de Carvalho, M.H. (2008). Drought stress and reactive oxygen species: production, scavenging and signaling. Plant Signaling & Behavior, 3, 156–165.

Daneshnia, F., Amini, A., & Chaichi, M. R. 2015. Surfactant effect on forage yield and water use efficiency for berseem clover and basil in intercropping and limited irrigation treatments. Agricultural Water Management, 160, 57–63.

Gao, Y., Duan, A., Sun, J., Li, F., Liu, Z., Liu, H., & Liu, Z. (2009). Crop coefficient and water-use efficiency of winter wheat/spring maize strip intercropping. Field Crops Research, 111, 65–73.

Grema, A.K., & Hess, T.M. (1994). Water balance and water use of millet-cowpea intercrops in north east Nigeria. Agricultural Water Management, 26, 169–185.

Hajiboland, R., Norouzi, F., & Poschenrieder, C. (2014). Growth, physiological, biochemical and ionic responses of pistachio seedlings to mild and high salinity. Trees, 28, 1065–1078.

Jan, S., Abbas, N., Ashraf, M., & Ahmad, P. (2019). Roles of potential plant hormones and transcription factors in controlling leaf senescence and drought tolerance. Protoplasma, 256, 313–329.

Koch, G., Rolland, G., Dauzat, M., Bédiée, A., Baldazzi, V., Bertin, N., Guédon, Y., & Granier, C. (2019). Leaf production and expansion: a generalized response to drought stresses from cells to whole leaf biomass– a case study in the tomato compound leaf. Plants, 8, 409.

Kong, C. H., Zhang, S. Z., Li, Y. H., Xia, Z. C., Yang, X. F., Meiners, S. J., & Wang, P. (2018). Plant neighbor detection and allelochemical response are driven by root-secreted signaling chemicals. Nature Communications, 9, 3867.

Krasensky, J., & Jonak, C. (2012). Drought, salt, and temperature stress-induced metabolic rearrangements and regulatory networks. Journal of Experimental Botany, 63, 1593–1608.

Kumar, S., Kaushal, N., Nayyar, H., & Gaur, P. (2012). Abscisic acid induces heat tolerance in chickpea (Cicer arietinum L.) seedlings by facilitated accumulation of osmoprotectants. Acta Physiologiae Plantarum, 34, 1651–1658.

Lemoine, R., Camera, S. L., Atanassova, R., Dédaldéchamp, F., Allario, T., Pourtau, N., Bonnemain, J. L., Laloi, M., Coutos-Thévenot, P., Maurousset, L., Faucher, M., Girousse, C., Lemonnier, P., Parrilla, J., & Durand, M. (2013). Source-to-sink transport of sugar and regulation by environmental factors. Frontiers in Plant Science, 4, 272.

Lugojan, C., & Ciulca, S. (2011). Evaluation of relative water content in winter wheat. Journal of Horticultural Science and Biotechnology, 15, 173–177.

Martin-Guay, M. O., Paquette, A., Dupras, J., & Rivest, D. (2018). The new green revolution: sustainable intensification of agriculture by intercropping. Science of the Total Environment, 615, 767–772.

Mc Adam, S. A., Brodribb, T. J., & Ross, J. J. (2016). Shoot‐derived abscisic acid promotes root growth. Plant, Cell Environment, 39, 652–659.

Mommer, L., Kirkegaard, J., & van Ruijven, J. (2016). Root–root interactions: towards a rhizosphere framework. Trends in Plant Science, 21, 209–217.

Noctor, G., Mhamdi, A., & Foyer, C. H. (2014). The roles of reactive oxygen metabolism in drought: not so cut and dried. Plant Physiology, 164, 1636–1648.

Parida, A. K., Dagaonkar, V. S., Phalak, M.S., & Aurangabadkar, L. P. (2008). Differential responses of the enzymes involved in proline biosynthesis and degradation in drought tolerant and sensitive cotton genotypes during drought stress and recovery. Acta Physiologiae Plantarum, 5, 619–627.

Rees, D. J. (1986). The effects of population density and intercropping with cowpea on the water use and growth of sorghum in semi-arid conditions in Botswana. Agricultural and Forest Meteorology, 37, 293–308.

Sadeghzadeh, N., Hajiboland, R., Moradtalab, N., Poschenrieder, C. 2021. Growth enhancement of Brassica napus under both deficient and adequate iron supply by intercropping with Hordeum vulgare: a hydroponic study. Plant Biosystems, 155, 632–646.

Sánchez-Blanco, M. J., Alvarez, S., Ortuño, M. F., & Ruiz-Sánchez, M. C. (2014). Root system response to drought and salinity: root distribution and water transport. In: Morte, A. & Varma, A. (Eds.) Root Engineering (pp 325–352). Germany, Springer.

Semchenko, M., Saar, S., & Lepik, A. (2014). Plant root exudates mediate neighbour recognition and trigger complex behavioural changes. New Phytologist, 204, 631–637.

Shackel, K. A., & Hall, A. E. (1984). Effect of intercropping on the water relations of sorghum and cowpea. Field Crops Research, 8, 381–387.

Singh, M., Kumar, J., Singh, S., Singh, V. P., & Prasad, S. M. (2015). Roles of osmoprotectants in improving salinity and drought tolerance in plants: a review. Reviews in Environmental Science and Biotechnology, 14, 407–426.

Singh, S., Narwal, S. S., & Chander, J. (1988). Effect of irrigation and cropping systems on consumptive use, water use efficiency and moisture extraction patterns of summer fodders. International Journal of Tropical Agriculture, 6, 76–82.

Skirycz, A., & Inze, D. (2010). More from less: plant growth under limited water. Current Opinion in Biotechnology, 21, 197–203.

Szabados, L., & Savouré, A. (2009). Proline: a multifunctional amino acid. Trends in Plant Science, 15, 89–97.

Tambussi, E. A., Bort, J., & Araus, J. L. (2007). Water use efficiency in C3 cereals under Mediterranean conditions: a review of physiological aspects. Annals of Applied Biology, 150, 307–321.

Tardieu, F., Simonneau, T., & Muller, B. (2018). The physiological basis of drought tolerance in crop plants: a scenario-dependent probabilistic approach. Annual Review of Plant Biology, 69, 733–759.

Tardieu, F., Granier, C., & Muller, B. (2011). Water deficit and growth. Coordinating processes without an orchestrator? Current Opinion in Plant Biology, 14, 283–289.

Verbruggen, N., & Hermans, C., (2008). Proline accumulation in plants: a review. Amino Acids, 35, 753–759.

Xiong, H., Shen, H., Zhang, L., Zhang, Y., Guo, X., Wang, P., Duan, P., Ji, C., Zhong, L., Zhang, F., & Zuo, Y. (2013). Comparative proteomic analysis for assessment of the ecological significance of maize and peanut intercropping. Journal of Proteomics, 78, 447–460.

Zuo, Y. (2013). Comparative proteomic analysis for assessment of the ecological significance of maize and peanut intercropping. Journal of Proteomics, 78, 447–460.

Xue, Y., Xia, H., Christie, P., Zhang, Z., Li, L., & Tang, C. (2016). Crop acquisition of phosphorus, iron and zinc from soil in cereal/legume intercropping systems: a critical review. Annals of Botany, 117, 363–377.

Zhou, S., Duursma, R. A., Medlyn, B. E., Kelly, J. W. G., & Prentice, I. C. (2013). How should we model plant responses to drought? An analysis of stomatal and non-stomatal responses to water stress. Agricultural and Forest Meteorology, 182/183, 204–214.

Zuo, Y., Li, X., Cao, Y., Zhang, F., & Christie, P. (2003). Iron nutrition of peanut enhanced by mixed cropping with maize: possible role of root morphology and rhizosphere microflora. Journal of Plant Nutrition, 26, 2093–2110.

Zuo, Y., Liu, Y., Zhang, F., & Christie, P. (2004). A study on the improvement iron nutrition of peanut intercropping with maize on nitrogen fixation at early stages of growth of peanut on a calcareous soil. Soil Science & Plant Nutrition, 50, 1071–1078.

Yemm, E. W., & Willis, A. J. (1954). The estimation of carbohydrates extracts by anthrone. Biochemistry Journal, 57, 508–514.

Fu, X., Wu, X., Zhou, X., Liu, S., Shen, Y., & Wu, F. (2015). Companion cropping with potato onion enhances the disease resistance of tomato against Verticillium dahliae. Frontiers in Plant Science, 6, 726.



20. 10. 2022



Original Scientific Article

How to Cite

SADEGHZADEH, N., HAJIBOLAND, R., & POSCHENRIEDER, C. (2022). Intercropping induces physiological and morphological plasticity in oilseed rape and barley under drought stress. Acta Agriculturae Slovenica, 118(3), 1–17.

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