Dissecting tissue- and species-specific responses of two Plantago species to waterlogging stress at physiological level
Introduction
Plantago L. is the largest genus of Plantaginaceae, and comprises about 270 species in the world, and including 20 in China. The aerial parts and seeds of several Plantago species have been widely used as drugs and food due to its considerable bioactivity (Beara et al., 2012, Li et al., 2009, Janković et al., 2012, Nishibe et al., 1995). Plantago asiatica L. (P. asiatica), a weed mainly distributed in temperate zone of Asia, has a wide altitudinal range of habitats, with the highest habitation at 2600 m elevation (Murai et al., 2009). The phytochemical components in P. asiatica have been surveyed by several groups (Nakaoki et al., 1961, Ravn et al., 1990, Nishibe et al., 1995). In contrast, Plantago fengdouensis (Z. E. Zhao & Y. Wang) Y. Wang & Z. Y. Li (P. fengdouensis), an endangered plant native to the Three Gorge Reservoir area of China, was discovered in only two small islands of two counties along the Yangtze River in the Three Gorges Reservoir of China in 2001, and identified and named in 2004 (Wang et al., 2004a, Wang et al., 2004b). Its natural habitat was seasonally flooded due to summer flooding of Yangtze River, and completely and perpetually submerged after the impoundment of the Three Gorges Reservoir (Wang et al., 2007).
Although a large number of Plantago species have been studied for the purpose of phytochemical analysis (Li et al., 2009, Janković et al., 2012, Nishibe et al., 1995, Nishibe, 2002), only few publications addressed the eco-physiological responses of Plantago species to abiotic stress. For example, the altitude variation had different effects on antioxidant system in leaves and in roots of P. major (Ren et al., 1999), and the ozone (O3) exposure resulted in a reduction in biomass and a decrease in the number of seeds production in P. major (Zheng et al., 2000). Under elevated CO2 condition, the biomass and total phenolic content of P. maritima increased, while protein content and the maximum carboxylation rate of Rubisco decreased (Davey et al., 2007). P. algarbiensis and P. almogravensis could efficiently accumulate aluminum from soil, especially in root tissues, and aluminum uptake was accompanied by substantial increases of citric, oxalic, malonic and fumaric acids contents in the plantlets of either species (Martins et al., 2013).
Waterlogging often occurs in many parts of the world every year. It often disturbs the plant growth, development, production, and many physiological functions, e.g., inhibition of photosynthesis and accumulation, increases of reactive oxygen species (ROS) production like superoxide radical (O2−), hydrogen peroxide (H2O2), and hydroxyl radical (OH), and occurrence of membrane lipid peroxidation (Alhdad et al., 2013, Candan and Tarhan, 2012, Yang et al., 2011). Accordingly, in order to cope with oxidative damages and membrane lipid peroxidation, plants possess a suite of antioxidant enzymatic system and produce some non-enzymatic components like glutathione and free proline to modulate the ROS levels (Alhdad et al., 2013, Candan and Tarhan, 2012, Erkan et al., 2008, Smirnoff and Cumbes, 1989, Yang et al., 2011, Yin et al., 2009c). Perennial plants inhabiting floodplains (or river islands) cannot escape from the flooding conditions and have to endure the occurrences of waterlogging (even flooding) stress. Accordingly, these plants often develop certain phenotypic plasticity and adaptive plasticity to cope with these stresses including adaptive adjustments in biomass allocation and life cycle (Chen and Xie, 2007, Pezeshki, 2001). The present study examined the hypothesis that P. fengdouensis originally inhabiting islands along the Yangtze River was more tolerant to waterlogging stress than P. asiatica widely distributing in Asia, especially in root systems. In addition, we also examined the hypothesis that tissue-specific responses to waterlogging stress between leaves and roots of Plantgo species. These analyses on physiological responses of P. fengdouensis to waterlogging stress may be used as indicators of reconstruction of the ex situ populations of the in situ conserved P. fengdouensis.
Section snippets
Plant materials and experimental design
The mature seeds of P. fengdouensis and P. asiatica were collected from their maternal plants grown in Wuhan Botanical Garden, Chinese Academy of Sciences in August, 2012. These seeds were used for germination in a greenhouse, which were maintained at 25–28 °C average temperature and 70–85% relative humidity. After germinating and growing for about 1 month, the seedlings were transplanted in pots (15 × 12 × 14 cm, length × width × height) filled with homogenized soil, one seedling per pot. After growth
Comparative analyses of morphological traits and the ratio of below-ground to above-ground biomass
As shown in Figs. 1A and 2A and C, P. fengdouensis had narrow leaves, whereas P. asiatica possessed wide leaves. On the other hand, P. fengdouensis developed stronger and deeper roots than P. asiatica did (Figs. 1A and 2B and D). In addition, P. fengdouensis had significantly higher ratio of below-ground biomass to above-ground biomass than P. asiatica under both control or stressed conditions (Fig. 1B). Waterlogging treatment inhibited the root development in both species, especially for the
Discussion
Most previous studies of Plantago species focused on their remarkable variety of curative properties, including astringent, styptic, antimicrobial, expectorant, diuretic and demulcent, and considerable bioactivity, including cytotoxic effects on cancer cell lines, anti-inflammatory, immuno-regulatory, antioxidant and antispasmodic effects in terms of biologically active compounds (Beara et al., 2012, Janković et al., 2012, Li et al., 2009, Nishibe et al., 1995). Only few studies addressed the
Acknowledgements
This work was sponsored by National Natural Science Foundation of China (No. 31270449), and “the Hundred Talents Program” and the Knowledge Innovation Project of Chinese Academy of Sciences (Grant nos. 54Y154761O01076 and 29Y329631O0263).
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