Yangjun Zou

Exogenous melatonin improves Malus resistance to Marssonina apple blotch

We examined whether exogenously applied melatonin could improve resistance to Marssonina apple blotch (Diplocarpon mali) by apple [Malus prunifolia (Willd.) Borkh. cv. Donghongguo]. This serious disease leads to premature defoliation in the main regions of apple production. When plants were pretreated with melatonin, resistance was increased in the leaves. We investigated the potential roles for melatonin in modulating levels of hydrogen peroxide (H2O2), as well the activities of antioxidant enzymes and pathogenesis‐related proteins during these plant–pathogen interactions. Pretreatment enabled plants to maintain intracellular H2O2 concentrations at steady‐state levels and enhance the activities of plant defence‐related enzymes, possibly improving disease resistance. Because melatonin is safe and beneficial to animals and humans, exogenous pretreatment might represent a promising cultivation strategy to protect plants against this pathogen infection.

Growth, biomass allocation, and water use efficiency of 31 apple cultivars grown under two water regimes

Plant growth, biomass allocation, carbon isotope composition (δ13C), and water use efficiency (WUE) of 31 cultivars of apple (Malus domestica Borkh.) grown under two water regimes were measured. Drought-stressed plants showed significant declines in tree height, trunk diameter, biomass production, and total leaf area, the extent to which depended upon cultivar. Also, gas exchange rates, instantaneous and long-term efficiencies (WUEI and WUEL, respectively), and values for δ13C differed among cultivars and watering regimes. Variations in WUEI were mainly due to changes in stomatal conductance (gs) under drought condition. ‘Qinguan’ and ‘Golden Delicious’ had greater trunk diameter, tree height, and had higher biomass production and WUEL under drought stress, implying that they are more suitable for arid and semi-arid regions. Moreover, WUEL was significantly and positively correlated with δ13C under two watering regimes, which suggests a potential for evaluating water use efficiency of Malus by measuring carbon isotope composition.

Genome-wide identification and expression profiling of the SnRK2 gene family in Malus prunifolia

Sucrose non-fermenting-1-related protein kinase 2 (SnRK2) constitutes a small plant-specific serine/threonine kinase family with essential roles in the abscisic acid (ABA) signal pathway and in responses to osmotic stress. Although a genome-wide analysis of this family has been conducted in some species, little is known about SnRK2 genes in apple (Malus domestica). We identified 14 putative sequences encoding 12 deduced SnRK2 proteins within the apple genome. Gene chromosomal location and synteny analysis of the apple SnRK2 genes indicated that tandem and segmental duplications have likely contributed to the expansion and evolution of these genes. All 12 full-length coding sequences were confirmed by cloning from Malus prunifolia. The gene structure and motif compositions of the apple SnRK2 genes were analyzed. Phylogenetic analysis showed that MpSnRK2s could be classified into four groups. Profiling of these genes presented differential patterns of expression in various tissues. Under stress conditions, transcript levels for some family members were up-regulated in the leaves in response to drought, salinity, or ABA treatments. This suggested their possible roles in plant response to abiotic stress. Our findings provide essential information about SnRK2 genes in apple and will contribute to further functional dissection of this gene family.

Influence of rootstock on antioxidant system in leaves and roots of young apple trees in response to drought stress

Grafting rootstocks are widely used to enhance plants resistance to various biologic and abiotic stresses. We determined how the rootstock genotype might influence plant responses to drought, using 2-year-old ‘Gale Gala’ apple trees grafted onto Malus sieversii and M. hupehensis. Under water stress, trees with the former as their rootstock had smaller reductions in rates of relative growth and photosynthesis, total biomass, leaf area, levels of leaf chlorophyll, and relative water content compared with those grafted onto the latter. They also had greater maximum photochemical efficiency and water-use efficiency. On the other hand, trees growing on M. sieversii rootstock had less production of superoxide radicals and hydrogen peroxide in both leaves and roots than those growing on M. hupehensis in response to drought stress. Furthermore, under drought conditions, leaves and roots from trees grafted onto M. sieversii had greater synthesis of ascorbic acid and glutathione, as well as higher activities of superoxide dismutase, catalase, ascorbate peroxidase, monodehydroascorbate reductase, dehydroascorbate reductase, and glutathione reductase. These results suggest that the choice of grafting rootstock can enhance drought resistance by improving the antioxidant system in a plant. Here, ‘Gale Gala’ trees grafted onto M. sieversii were more drought-resistant than those on M. hupehensis rootstock.

Interactive effects of water and nitrogen supply on growth, biomass partitioning, and water-use efficiency of young apple trees

Drought and low concentration of soil nitrogen (N) limit crop growth and production in arid and semiarid regions. Improving the efficient use of these limited resources is an important challenge. We tested one-year-old trees of three apple cultivars (Malus domestica cv. ‘Golden Delicious’, ‘Naganofuji No. 2’, and ‘Pink Lady’) grafted on M. hupehensis to determine how water and N supplies influence growth, biomass production, and water-use efficiency (WUE). Two watering regimes (80 and 50% field water capacity) and three N supplies (N0: 0, NL: 75, and NH: 150 mg N kg −1 soil) were used. Drought dramatically diminished plant height (PH), basal diameter (BD), biomass production, total leaf area (LA), specific leaf area (SLA), leaf relative water content (RWC), photosynthetic rate (Pn), stomatal conductance (gs), and WUE, but increased root/shoot ratio. NL significantly increased PH, BD, LA, SLA, RWC, Pn, gs, and WUE, but NH reduced or had little influence on these parameters. Pn, gs, LA, and root biomass significantly and positively correlated with WUE. Our results suggest that water and N are corelated, and NL enhance drought tolerance and WUE through increased photosynthetic capacity and water uptake. Thus, an appropriately low N supply would be recommended under dry condition, but excess N supply should be avoided.

Growth, gas exchange, water-use efficiency, and carbon isotope composition of ‘Gale Gala’ apple trees grafted onto 9 wild Chinese rootstocks in response to drought stress

To determine the effects of rootstock choice on the scion response to drought stress, we compared the vegetative growth, biomass accumulation, gas exchange, and water-use efficiency (WUE) of ‘Gale Gala’ apple (Malus domestica Borkh.) trees grafted onto nine wild Chinese Malus rootstocks. Compared with the well-watered control, drought treatment limited growth, as manifested by smaller increments in plant height (PH), trunk diameter (TD), total fresh biomass (TB), total dry biomass (TDB), total leaf area (LA), and relative growth rate (RGR). The extent of this effect differed among rootstocks. Stress conditions led to increases in the root/shoot ratio (RSR), leaf thickness (LT), water-holding capacity (WHC), carbon isotope composition (δ13C), and WUE. Decreases were noted in stomatal density (SD), leaf relative water content (RWC), chlorophyll content (Chl), net photosynthetic rate (PN), transpiration rate (E), and stomatal conductance (gs), again varying by rootstock. Those that are generally considered more drought-tolerant, e.g., M. sieversii, M. prunifolia, and M. toringoides, had smaller declines in PH, TD, TB, TDB, LA, RGR, SD, RWC, Chl, PN, E, and gs and proportionally greater increases in RSR, LT, WHC, δ13C, and WUE compared with the droughtsensitive M. hupehensis and M. sieboldii. These results suggest that moisture stress has a significant dwarfing effect in the latter two species. Based on WUE calculations, trees on drought-tolerant rootstocks showed higher tolerance when stressed, whereas those on drought-sensitive rootstocks were less tolerant, as indicated by their lower WUE values.

Responses of young ‘Pink lady’ apple to alternate deficit irrigation following long-term drought: growth, photosynthetic capacity, water-use efficiency, and sap flow

We studied photosynthetic capacity, growth, sap flow, and water-use efficiency in young trees of ‘Pink Lady’ apple (Malus domestica) that were exposed to 60 d of moisture stress. Three irrigation schemes were tested in the greenhouse: well-watered control; drought; or alternate deficit irrigation (ADI). Compared with the drought-stressed plants, those treated via ADI showed better height growth, larger scion diameters, and greater total leaf area, as well as significantly increased gains in dry biomass and rootstock diameters. However, their performance was still significantly lower than that demonstrated by continuously well-watered plants. Sap flow was greater under ADI than under drought, but less than under control conditions. The average rate of net photosynthesis, total amount of irrigation water applied, and dry biomass gain had highly significant and positive linear correlations with long-term water-use efficiency (WUEL). The same was true between average stomatal conductance and WUEL. By contrast, instantaneous water-use efficiency (WUEI) was very significantly and negatively correlated with WUEL. In addition, values for WUEL were much higher from well-watered plants when compared with either drought-stressed trees or those treated per ADI. Therefore, our results indicate that, although ‘Pink Lady’ apple normally has high WUE, it still consumes a large amount of water. Therefore, the practice of ADI following a period of long-term drought could be used to improve growth and WUEL by this cultivar.

Genome-wide identification of genes involved in polyamine biosynthesis and the role of exogenous polyamines in Malus hupehensis Rehd. under alkaline stress

Polyamines (PAs) in plants are growth substrates with functions similar to phytohormones. Although they contribute to diverse processes, little is known about their role in stress responses, especially for perennial woody plants. We conducted a genome-wide investigation of 18 sequences involved in PA biosynthesis in the genome of apple (Malus domestica). Further analysis was performed to construct a phylogenetic tree, analyze their protein motifs and gene structures. In addition, we developed their expression profiles in response to stressed conditions. Both MDP0000171041 (MdSAMDC1) and MDP0000198590 (MdSPDS1) were induced by alkaline, salt, ABA, cold, and dehydration stress treatments, suggesting that these genes are the main contributors to activities of S-adenosylmethionine decarboxylase (EC 4.1.1.50) and spermidine synthase (EC 2.5.1.16) in apple. Changes in PA biosynthesis under stress conditions indicated that spermidine and spermine are more essential than putrescine for apple, especially when responding to alkaline or salt stress. When seedlings of M. hupehensis Rehd. were supplied with exogenous PAs, their leaves showed less chlorosis under alkaline stress when compared with untreated plants. This application also inhibited the decline in SPAD levels and reduced relative electrolyte leakage in those stressed seedlings, while increasing their concentration of active iron. These results suggest that the alteration in PA biosynthesis confers enhanced tolerance to alkaline stress in M. hupehensis Rehd.