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Effects of Water and Energy on Plant Diversity Along the Aridity Gradient Across Dryland in China

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Effects of Water and Energy on Plant Diversity Along the Aridity Gradient Across Dryland in China plants Article Effects of Water and Energy on Plant Diversity along the Aridity Gradient across Dryland in China Shuran Yao 1,†, Muhammad Adnan Akram 1,† , Weigang Hu 1 , Yuan Sun 1 , Ying Sun 1, Yan Deng 1,2, Jinzhi Ran 1,* and Jianming Deng 1 1 State Key Laboratory of Grassland Agro-Ecosystem, School of Life Sciences, Lanzhou University, Lanzhou 730000, China; [email protected] (S.Y.); [email protected] (M.A.A.); [email protected] (W.H.); [email protected] (Y.S.); [email protected] (Y.S.); [email protected] (Y.D.); [email protected] (J.D.) 2 State College of Forestry, Southwest Forestry University, Bailongsi 300, Kunming 650224, China * Correspondence: [email protected] † These authors contributed equally to this work. Abstract: Plants need water and energy for their growth and reproduction. However, how water and energy availability influence dryland plant diversity along the aridity gradient in water-limited re- gions is still lacking. Hence, quantitative analyses were conducted to evaluate the relative importance of water and energy to dryland plant diversity based on 1039 quadrats across 184 sites in China’s dryland. The results indicated that water availability and the water–energy interaction were pivotal to plant diversity in the entire dryland and consistent with the predictions of the water–energy dynamic hypothesis. The predominance of water limitation on dryland plant diversity showed a weak trend with decreasing aridity, while the effects of energy on plants were found to be significant in mesic regions. Moreover, the responses of different plant lifeforms to water and energy were found to vary along the aridity gradient. In conclusion, the study will enrich the limited knowledge Citation: Yao, S.; Akram, M.A.; Hu, W.; Sun, Y.; Sun, Y.; Deng, Y.; about the effects of water and energy on plant diversity (overall plants and different lifeforms) in the Ran, J.; Deng, J. Effects of Water and dryland of China along the aridity gradient. Energy on Plant Diversity along the Aridity Gradient across Dryland in Keywords: aridity gradient; dryland; plant diversity; plant lifeforms; water–energy dynamics China. Plants 2021, 10, 636. https:// doi.org/10.3390/plants10040636 Academic Editor: Daniel Sánchez-Mata 1. Introduction The growth and reproduction of plants are dependent on water availability and envi- Received: 18 February 2021 ronmental energy [1,2]. Environmental energy (heat) is the basic requirement of all plants Accepted: 22 March 2021 needed to perform the different functions and activities of their lives, such as photosynthe- Published: 27 March 2021 sis. Water is an integral part of plant tissues and without it, plants cannot perform many physiological activities [3,4]. Environmental energy and water are interrelated [3]; for ex- Publisher’s Note: MDPI stays neutral ample, energy can increase the water-use efficiency of plants to promote photosynthesis, with regard to jurisdictional claims in as snow can reduce the freezing effect on plants [1,5]. However, excessively high or low published maps and institutional affil- energy may change the water into vapor or ice, and these physical processes could result iations. in the limitation of a plant’s growth and development [6,7]. In contrast, when energy is abundant, water restriction (e.g., limiting water metabolic processes and closing of the stomata) also prevents plants from using the energy for photosynthesis [8–10]. The effects of water and energy on constraining the physicochemical and physio- Copyright: © 2021 by the authors. logical processes of plants might generate and maintain a broad gradient of plant diver- Licensee MDPI, Basel, Switzerland. sity [1,6,11–13]. The water–energy dynamic hypothesis emphasizes that the large-scale This article is an open access article gradient of species diversity is related to both water and energy [11,13–15]. It provides distributed under the terms and theoretical support to understand the environmental capacity to support greater plant- conditions of the Creative Commons species richness [6,16]. The relative importance of water and energy maintenance to plant Attribution (CC BY) license (https:// creativecommons.org/licenses/by/ diversity may depend on climatic regions [17,18]. For example, water is a limiting factor 4.0/). in warm regions, where the energy is sufficient; while energy becomes more important Plants 2021, 10, 636. https://doi.org/10.3390/plants10040636 https://www.mdpi.com/journal/plants Plants 2021, 10, x FOR PEER REVIEW 2 of 16 Plants 2021, 10, 636 2 of 15 to plant diversity may depend on climatic regions [17,18]. For example, water is a limit- ing factor in warm regions, where the energy is sufficient; while energy becomes more important in cold regions, where inputs of energy are low [15]. Conversely, the rich- in cold regions, where inputs of energy are low [15]. Conversely, the richness–energy ness–energy relationship depends on water availability [6]. Species diversity also has relationship depends on water availability [6]. Species diversity also has shown the strong shown the strong effects of energy and water on woody plant species [1]. Nevertheless, effects of energy and water on woody plant species [1]. Nevertheless, how water and how water and energy availability interact to influence dryland plant diversity in local energy availability interact to influence dryland plant diversity in local communities along communities along the water gradient in water-limited regions is still an open question. the water gradient in water-limited regions is still an open question. The United Nations Environmental Program (UNEP) defines drylands as arid re- The United Nations Environmental Program (UNEP) defines drylands as arid regions gions where the aridity index (the ratio of annual precipitation to annual potential evap- where the aridity index (the ratio of annual precipitation to annual potential evapotran- otranspiration) is less than 0.65 [19]. Based on the aridity index, drylands can be classified spiration) is less than 0.65 [19]. Based on the aridity index, drylands can be classified as dry-subhumid regions, semiarid regions, arid regions, and hyperarid regions [20,21]. as dry-subhumid regions, semiarid regions, arid regions, and hyperarid regions [20,21]. ApproximatelyApproximately 45% 45% of of Earth’s Earth’s land-surface land-surface area area is aridis arid [22 [22].]. In China’sIn China’s dryland, dryland, ephemeral ephem- plants,eral plants, annual annual herbs, perennialherbs, perennial herbs, and herb woodys, and plants woody (including plants shrubs,(including sub-shrubs, shrubs, andsub-shrubs, small trees) and are small the dominanttrees) are the plant dominant lifeforms, plant but talllifeforms, trees are but rare. tall Differenttrees are lifeformsrare. Dif- haveferent distinct lifeforms physiological have distinct attributes physiological and growth attributes forms [23and–25 growth]. For example, forms [23–25]. ephemeral For plantsexample, are ephemeral a special group plants of are dryland a special herbaceous group of plantsdryland that herbaceous typically dependplants that on tran-typi- sientcally precipitationdepend on transient and temperature, precipitation consequently and temperature, growing consequently fast and adapting growing to thefast dryand environmentadapting to the [23 ,dry24]. environment Shrubs have a[23,24]. strong Shru rootingbs have system a strong to effectively rooting extract system moisture to effec- intively low-moisture extract moisture environments in low-moisture [2,26]. Different environments lifeforms [2,26]. have Different distinct adaptive lifeforms strate- have giesdistinct to survive adaptive in severestrategies environmental to survive conditionsin severe [environmental25], which may conditions lead to their [25], different which responsesmay lead to watertheir different and energy responses along the to water aridity and gradient. energy along the aridity gradient. InIn thisthis study, study, 184 184 sites sites were were surveyed surveyed and and 1039 1039 quadrats quadrats were were sampled sampled along along the arid- the ityaridity gradient gradient in the entirein the dryland entire (includingdryland hyperarid,(including arid,hyperarid, semiarid, arid, and dry-subhumidsemiarid, and regions)dry-subhumid of China. regions) The of objectives China. The of theobjectives present of studythe present were: study (1) to were: assess 1) theto assess relative the importancerelative importance of water of and water energy and energy to overall to overall plant diversity plant diversity along along the aridity the aridity gradient; gra- anddient; (2) and to evaluate 2) to evaluate whether whether the effects the ofeffect waters of and water energy and on energy plant diversityon plant ofdiversity different of lifeformsdifferent varylifeforms along vary the along aridity the gradient. aridity gradient. In the light In the of distinctlight of adaptivedistinct adaptive strategies strat- of differentegies of different plant lifeforms, plant lifeforms, it is predicted it is pred thaticted water that and water energy and might energy influence might influence the lifeform the diversitylifeform diversity differently. differently. 2.2. MaterialsMaterials andand MethodsMethods 2.1.2.1. StudyStudy RegionRegion The region is located in the northwestern dryland of China (36.01◦~50.70◦ N and ◦The region◦ is located in the northwestern dryland of China (36.01°~50.70° N and 76.6276.62°~122.40°~122.40 E) and occupies approximately 2.442.44 millionmillion squaresquare
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