Chenopodium quinoa Willd. is an annual facultative halophytic pseudocereal widely studied for its physiology and grain yield owing to its great tolerance to unfavorable growing conditions. However, the morphophysiological and anatomical characteristics of plants’ photosynthetic organs under various and combined abiotic stresses during the early stages of development have not been thoroughly studied. Therefore, the current study compared the influence of osmotic, salt, and combined stress at different intensities on the morphology and anatomy of photosynthetic organs in young quinoa plants. The main findings demonstrate that salt stress at an intensity between 100 and 200 mM NaCl is not critical for the growth of young quinoa plants and that the young plants can withstand salt stress at an intensity of 300 mM NaCl. However, it can be concluded that some adaptation mechanisms of the plants were already violated at a salt stress intensity of 200 mM NaCl, while significant changes in the water balance of the plants were observed at an intensity of 300 mM NaCl, possibly caused by damage to the cell structures.

Chenopodium quinoa is a genetically diverse crop that can adapt to a wide range of environments, including temperatures and salinities. However, only a few studies have assessed the combined effects of two or more environmental factors on C. quinoa. Here, we investigated the effects of salinity (300 mM NaCl), elevated temperature (35 °C), and their interaction with growth, water–salt balance, the efficiency of photosystem II (PSII), the activity of cyclic electron transport (CET) around photosystem I (PSI), Rubisco and PEPC enzyme content, and the expression of photosynthetic genes. We found that elevated temperature did not decrease the biomass but caused a significant increase in the water and potassium content of C. quinoa leaves. The decrease in PSII efficiency under elevated temperature was accompanied by an increase in the expression of genes encoding the components of PSII (psbA) and linear electron transport (FDI), as well as the main photosynthetic protein Rubisco (rbcL). Moreover, the strongest effect was induced by the combined effect of elevated temperature and salinity, which induced high oxidative stress (a threefold increase in MDA), a threefold decrease in the biomass, a twofold decrease in PSII efficiency, and a two- to eightfold decrease in the expression of the photosynthetic genes psbA, FDI, and rbcL. PSI was more tolerant to all forms of stress; however, the combined effect of elevated temperature and salinity downregulated the expression of PGR5 and FNR1, which may diminish the role of PGR5/PGRL1-dependent CET in favor of the NDH-dependent CET of PSI. The obtained data on the functioning of photosystems and the expression of photosynthetic genes under combined stress (elevated temperature and salinity) can make a significant contribution to understanding the mechanisms of tolerance of C. quinoa to multiple stresses under climate change conditions.

Plant adaptation to salinity is a highly multifaceted process, harnessing various physiological mechanisms depending on the severity and duration of salt stress. This study focuses on the effects of 4- and 10-day treatments with low (100 mM NaCl) and moderate (200 mM NaCl) salinity on growth, CO2/H2O gas exchange, stomatal apparatus performance, the efficiency of photosystems I and II (PS I and II), content of key C4 photosynthesis enzymes, and the accumulation of Na+, K+, and proline in shoots of the widespread forage C4 halophyte Kochia prostrata. Our data show that 4 days of low salinity treatment resulted in a decrease in biomass, intensity of apparent photosynthesis, and cyclic electron transport around PS I. It was accompanied by an increase in transpiration and Rubisco and PEPC contents, while the Na+ and proline contents were low in K. prostrata shoots. By the 10th day of salinity, Na+ and proline have accumulated; PS I function has stabilized, while PS II efficiency has decreased due to the enhanced non-photochemical quenching of chlorophyll fluorescence (NPQ). Thus, under low salinity conditions, Na+ accumulated slowly and the imbalance between light and dark reactions of photosynthesis was observed. These processes might be induced by an early sodium signaling wave that affects cellular pH and ion homeostasis, ultimately disturbing photosynthetic electron transport. Another adaptive reaction more “typical” of salt-tolerant species was observed at 200 mM NaCl treatment. It proceeds in two stages. First, during the first 4 days, dry biomass and apparent photosynthesis decrease, whereas stomata sensitivity and dissipation energy during dark respiration increase. In parallel, an active Na+ accumulation and a decreased K+/Na+ ratio take place. Second, by the 10th day, a fully-fledged adaptive response was formed, when growth and apparent photosynthesis stabilized and stomata closed. Decreased dissipation energy, increased WUE, stabilization of Rubisco and PEPC contents, and decreased proline content testify to the completion of the adaptation and stabilization of the physiological state of plants. The obtained results allowed us to conclude that the formation of a full-fledged salt-tolerant response common for halophytes in K. prostrata occurs by the 10th day of moderate salinity.

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