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Comparison of Physiological and Antioxidant Responses of Anoda Cristata and Cotton to Progressive Drought

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Comparison of Physiological and Antioxidant Responses of Anoda Cristata and Cotton to Progressive Drought DOI: 10.1111/j.1365-3180.2012.00924.x Comparison of physiological and antioxidant responses of Anoda cristata and cotton to progressive drought HH RATNAYAKA*, WT MOLIN &TMSTERLINGà *Department of Biology, Xavier University of Louisiana, New Orleans, LA, USA, Southern Weed Science Research Unit, Stoneville, MS, USA, and àLand Resources and Environmental Sciences, Montana State University, Bozeman, MT, USA Received 4 August 2011 Revised version accepted 3 April 2012 Subject Editor: Frank Forcella, USDA-ARS, USA and F , but increased xanthophyll cycle conversion Summary PSII state in both A. cristata and cotton. Anoda cristata had Knowledge about physiological responses of weed and prolonged gas exchange at lower leaf water contents crop to drought is needed to minimise crop losses caused than cotton. Air-to-leaf temperature difference was at by weed interference and to improve cropping systems least 66% greater in A. cristata, while a-tocopherol and crop cultivars. In this study, gas exchange, photo- concentration increased only in cotton under progressive chemistry and antioxidant defences of two Anoda drought. Anoda cristata appears to use patchy stomatal cristata (spurred anoda) accessions from Mississippi closure and alternative electron sinks to minimise light and New Mexico and two cotton types (Gossypium damage during severe drought. Furthermore, sustained hirsutum and Gossypium barbadense) were evaluated water-holding capacity and assimilation helped by under unstressed conditions and progressive drought. prolonged stomatal activity, larger pigment pools and Under no stress, net photosynthesis (Pnet) and quantum lower leaf temperature in A. cristata may confer stress yield (FPSII) were similar, but stomatal conductance (gs) tolerance and consequent success in cotton – A. cristata was higher in A. cristata than cotton. Glutathione interference. reductase activity was highest in A. cristata from Mis- Keywords: spurred anoda, photosynthesis, gas exchange, sissippi. Cotton had at least 92% greater a-tocopherol photochemistry, stress tolerance, oxidative stress, stoma- concentration than A. cristata. Progressive drought that tal activity, leaf temperature. suppressed gas exchange to near zero decreased biomass RATNAYAKA HH, MOLIN WT & STERLING TM (2012). Comparison of physiological and antioxidant responses of Anoda cristata and cotton to progressive drought. Weed Research 52, 358–366. Introduction a serious problem in other crops ⁄ countries (Marzocca, 1976; Puricelli et al., 2003). Furthermore, there are Anoda cristata (L.) Schlecht. (spurred anoda) was once regions where herbicide applications are not feasible, competitive and difficult to manage in cotton (Gossypium because of reasons such as economics, restrictions and hirsutum L. and Gossypium barbadense L.) (Arle & resistance concerns, especially in weeds with persistent Hamilton, 1973; Patterson et al., 1988) and was listed seed banks as with A. cristata (Puricelli et al., 2002). among the 10 most important cotton weeds in the United Thus, studies on mechanisms of stress tolerance of weed States (Dowler, 1992). While the use of herbicides, and crop continue to be important in the development especially glyphosate, has lessened its impact and occur- of cropping systems and crop cultivars that minimise rence in most cotton farms in the United States, it is still losses because of weeds without the use of herbicides. Correspondence: HH Ratnayaka, Department of Biology, Xavier University of Louisiana, 1 Drexel Drive, New Orleans, LA 70125, USA. Tel: (+1) 504 520 5709; Fax: (+1) 504 520 7918; E-mail: [email protected] Ó 2012 The Authors Weed Research Ó 2012 European Weed Research Society Weed Research 52, 358–366 Mechanisms of drought tolerance 359 Reduced productivity in cotton under A. cristata and rinsed with water for 30 min before germination. interference is in part due to deprivation of water, Anoda cristata and cotton (Delta and Pine Land cv. 5415 caused by interspecific rather than intraspecific interfer- [G. hirsutum] and cv. Pima S-7 [G. barbadense]) seeds ence (Ratnayaka et al., 2003). Like cotton, A. cristata is were germinated on wet paper towels. Seedlings were in the Malvaceae and has a similar growth habit. first planted into 120-mL pots containing Terra-Lite Moreover, it has several clearly identifiable biotypes ⁄ Metro Mix 360 (W. R. Grace & Co., Memphis, TN, accessions. Thus, it offers a promising model to under- USA). Plants with the emerging first true leaf were stand how drought stress is initiated and tolerated at the transplanted into 8-L pots containing the same medium physiological level in cotton–weed interactions. Insight and grown in the glasshouse under natural day length in into photochemical or stomatal limitations to net spring. Twelve plants of each A. cristata accession were assimilation during different stages of a progressive maintained per each of six replications arranged in a drought may help identify means to mitigate these stress randomised complete block design. Temperature was effects. Furthermore, because cultivars of cotton devel- maintained at c. 30 and 18°C during day and night, oped with minimum stomatal limitations to photosyn- respectively, and relative humidity was between 20 and thesis may be suitable only for irrigated cropping 40%. Plants were watered daily and fertilised weekly systems, biochemical limitations may represent effective with Technigro 20-18-18 fertiliser (Fisons Horticulture foci when searching for drought-resistant genotypes Inc., Warwick, NY, USA) until the progressive drought (Hutmacher & Krieg, 1983). Thus, clues for developing was imposed at 7 weeks after planting. Watering was stress-resistant crops may involve assessments of prop- withheld from six randomly selected plants of a given erties such as non-photochemical energy dissipation of plant type in each replication, while the other six plants PSII, antioxidants such as a-tocopherol and b-carotene, served as well-watered controls (watered uniformly over and enzymes that regulate the redox buffer system, such the pot to field capacity until dripping). Measurements as catalase, ascorbate peroxidase and glutathione reduc- were taken from a randomly selected water-stressed and tase (GR) (Kunert & Foyer, 1993; Secenji et al., 2008), control plant of a given plant type in each replication on under both stressed and unstressed conditions, along each of the 6 days between drought initiation and wilt. with stomatal regulation. The experiment was repeated in time. Assimilation, PSII activity and antioxidant systems of A. cristata accessions from arid and humid regions Gas exchange, leaf temperature and quantum yield have not been compared under irrigated conditions and progressive drought. The goal of this study was to Gas exchange variables and leaf temperature of the identify physiological mechanisms of drought tolerance fourth leaf from the apex was recorded with an infrared in both weed and cotton types, to help develop cropping gas analyser-based photosynthesis system (LI-6400, LI- systems strategies and drought- and weed-tolerant COR Inc. Lincoln, NE, USA). All measurements were cotton cultivars. Thus, the hypotheses of this study are taken after net photosynthesis (Pnet) and stomatal (i) constitutive levels of gas exchange, quantum yield conductance (gs) stabilised on the graphical display of and antioxidant systems of two A. cristata accessions the instrumentÕs console. Measurement conditions were ) ) and two cotton cultivars are different, and (ii) these four as follows: internal photon flux density 500 lmol m 2 s 1 plant types will experience different limitations to to match the average light level at plant height in the )1 assimilation and PSII activity and resort to different glasshouse, flow rate 400 lmol s and internal CO2 )1 components of protective antioxidant systems during concentration 400 lmol mol . Quantum yield (FPSII) progressive drought. Changes in leaf bulk water status, was measured using the same leaf with OS5-FL relative water content (RWC), was used as the reference modulated chlorophyll fluorometer (Opti-Sciences Inc., 0 for the degree of drought. Tyngsboro, MA, USA). Minimal fluorescence (Fo) was measured after a 3 s pulse of far-red light, and maximal 0 fluorescence (Fm) was measured after a saturation pulse Materials and methods ) ) of 0.8 s at 5000 lmol m 2 s 1. The measuring beam was hiset at 360 lmol m)2 s)1. Quantum yield was computed as Plant material and drought treatment 0 0 0 0 Fm À Ft =Fm , where Ft is the fluorescence immedi- Seeds of New Mexico (NM, arid) and Mississippi (MS, ately before saturation pulse. humid) accessions of A. cristata were collected from Leyendecker Research Farm of New Mexico State Antioxidant enzyme activities and pigments University and USDA-ARS, Stoneville, MS, respec- tively. They were scarified by immersion in concentrated Leaf discs (0.6 cm diam) were taken using a paper hole- sulphuric acid (9.3 N) for 20 min (Solano et al., 1976) puncher from the same leaf used to measure Pnet. Discs Ó 2012 The Authors Weed Research Ó 2012 European Weed Research Society Weed Research 52, 358–366 360 HH Ratnayaka et al. were frozen immediately on dry ice and stored at –20°C 40 until used for biochemical assays. Ten leaf discs were Water-stressed Unstressed homogenised in 700 lL of a solution containing 50 mM * 30 PIPES buffer, 6 mM cysteine hydrochloride, 10 mM *** D-isoascorbic acid, 1 mM EDTA, 1% polyvinylpyrroli- done (PVP-10), 0.3% triton X-100, pH 6.8, and centri- 20 fuged at 7000 g for 15 min at 4°C, as modified from ns Foster and Hess (1980). The supernatant was desalted Shoot biomass (g) and used for assaying total protein and enzyme activities 10 * (APX, ascorbate peroxidase; CAT, catalase; GR, gluta- thione reductase), as described previously (Bettmann 0 et al., 2006). Antioxidants and chlorophylls were MS NM DP Pima assayed according to a method modified from Gilmore Fig. 1 Shoot biomass of the two Anoda cristata accessions and Yamamoto (1991), as described previously (Bett- (Mississippi and New Mexico) and two cotton cultivars (Delta Pine mann et al., 2006). Xanthophyll cycle conversion state 5414 and Pima S-7) at the end of experiments. n = 36.
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