Effects of post–silking low temperature on the physicochemical properties of waxy maize starch
Introduction
Low temperature (LT) is an important abiotic stress that seriously affects plant growth and development and influences crop productivity and quality [1], [2], [3], [4]. LT considerably affects enzymatic activities related to starch biosynthesis and enhances granule bound starch synthase I [1]; therefore, it increases the amylose–amylopectin ratio [5]. Additionally, amylopectin chain–length is increased by the low activities of starch synthase and starch branching enzyme, which are responsible for amylopectin branching and elongation [5], [6], [7]. LT decreases the activities of starch synthase, sucrose synthase, and invertase, which results in a decrease in the contents of sucrose and many amino acids in chickpea [8]. LT also induces hormonal imbalance, reduces carbohydrate supply, increases grain abortion, and decreases grain filling rate [9]. Changes in starch metabolic enzymes and hormones affect grain quality [10]. LT during grain filling reduces peak, trough, and final viscosities [11], as well as the gelatinization temperatures, enthalpy, and deteriorate the grain quality of rice [12]. LT declines grain appearance and milling and cooking quality traits but enhances chalkiness, broken rice, and gelatinization temperature; the most responsive period of rice quality to extreme temperatures occurs during the second week of post–heading [13]. LT increases the number of mid–sized and large granules with irregular shape and reduces the relative crystallinity (RC) of rice grains [12]. Labuschagne et al. [14] suggested that LT at the grain–filling stage reduces the total starch content and increases the amylose content in different wheat cultivars. LT increases 2–AP biosynthesis, cooked rice elongation percentage, and amylose content [15]. Aoki et al. [16] reported that bread firming is slowed in rice flour filled at LT. In general, LT has a positive effect on the protein concentrations and processing quality of wheat but has a negative effect on starch concentration and appearance quality [17].
High temperature is a normal climatic condition during summer and affects maize plant growth, development, yield, and quality. Adjusting the sowing date can improve maize adaption to climate change; optimal sowing date can lead to high yield and good quality [18]. In China, the postponement of the sowing date for summer maize is a simple and efficient cultivation option to escape or avoid heat stress, but this method increases the probability of cold stress [19]. Study on maize in North China Plain reported that grain yield was affected by the sowing dates (mid–March to mid–July) and that plants suffered from high temperature (>30.2 °C) and LT (<20.7 °C) during grain filling under early and late sowing conditions, respectively, and ultimately decreased grain weight [20]. The sowing date of summer maize ranges from late May to early August in Southern China. Sowing early or late can induce heat or cold stress at later growth stages, respectively. Previous study focused on the effects of heat stress on the physicochemical properties of maize starch [21]. In comparison to normal temperature (AT, 25 °C), LT (16 °C) during grain filling disturbed hormone balance, decreased enzymatic activities related to starch biosynthesis during the mid–grain filling period, and limited starch accumulation [10]. However, the effect of LT stress on the physicochemical properties of maize remains unknown. The starch of waxy maize, a special maize type, is composed of nearly pure amylopectin, which endows high viscosity, high transmittance, high stability, and low retrogradation [22]. The starch physicochemical properties of waxy maize are affected by sowing dates [23], [24]. We hypothesized that post–silking LT affects the structural and functional properties of waxy maize starch. We verified this hypothesis by estimating the physicochemical properties of waxy maize starch in response to LT at the late growth stage. The results can provide a basis for the efficient utilization of waxy maize starch in areas that suffer LT at the late growth stage.
Section snippets
Experimental design
Two waxy maize hybrids, namely 'Suyunuo5' (SYN5) and 'Yunuo7' (YN7), which are widely planted in Southern China, were used as experimental materials.
Pot (2019) and field trials (2018–2019) were conducted at the Experimental Farm of Yangzhou University (Yangzhou, China). For the pot trial, plastic pots with 38 cm height and 43 cm diameter were loaded with 30 kg sieved sandy loam soil. Three seeds were sown in each pot in 1 July, and one plant was retained at the six–leaf stage. The plants were
Morphology of starch granules
Starch exists as granules in amyloplasts, and the granules in the endosperm increase in number in the first two weeks after pollination and subsequently enlarge in size [26]. In the field trial, most starch granules under AT condition presented abnormal or irregular polytopes, whereas more small granules with oval shapes were observed under LT condition in both hybrids in both years (Fig. 1). The granule surface under both AT and LT was smooth compared with the rough granules under high
Conclusion
The morphological, structural, and functional properties of waxy maize starch were remarkably affected by growth temperatures at the late growth stages. The plants that suffered post–silking LT stress in pot and field trials produced starch with more small granules and oval or round shapes in both hybrids in both years. The small SGS, low RC, high proportion of short chains, and short amylopectin chain length of starch under LT conditions endow low pasting viscosities, gelatinization enthalpy,
CRediT authorship contribution statement
Huan Yang: Investigation, Formal analysis, Writing–original draft, Writing–review & editing. Qi Wei: Investigation, Formal analysis, Writing–review & editing. Weiping Lu: Conceptualization, Methodology, Supervision. Dalei Lu: Conceptualization, Funding acquisition, Methodology, Supervision, Validation, Writing–review & editing.
Declaration of competing interest
The authors declare that they have no conflicts of interest.
Acknowledgement
This study was supported by the National Natural Science Foundation of China (31771709, 32071958), Jiangsu Agriculture Science and Technology Innovation Fund (CX[20]3147), Priority Academic Program Development of Jiangsu Higher Education Institutions, and High–end Talent Support Program of Yangzhou University.
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The authors contribute equally to this work.