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Crop Science 41:133-138 (2001)
© 2001 Crop Science Society of America

TURFGRASS SCIENCE

Zoysiagrass Water Relations

R.H. Whitea, M.C. Engelkeb, S.J. Andersonb, B.A. Ruemmelec, K.B. Marcumd and G.R. Taylor, IIa

a II, Soil & Crop Sciences Dep., Texas A&M Univ., College Station, TX 77843-2474
b Texas A&M Research and Extension Center, Dallas, TX 75252
c Univ. of Rhode Island, Kingston, RI
d Dep. of Plant Sciences, Univ. of Arizona, Tucson, AZ

Corresponding author (rh-white{at}tamu.edu)


   ABSTRACT
TOP
 NOTES
 ABSTRACT
INTRODUCTION
 MATERIALS AND METHODS
 RESULTS AND DISCUSSION
 REFERENCES
 
Development of drought resistant, water-conserving cultivars continues to be an objective of turfgrass breeding programs. This study was conducted to determine for 15 zoysiagrasses [Zoysia japonica Steud., Z. matrella (L.) Merr., and Z. japonica Steud. x Z. tenuifolia Willd.] under greenhouse conditions (i) water relations characteristics, (ii) survival and recovery from extreme water stress, and (iii) the relationship of water relations characteristics to supplemental irrigation requirement determined in field studies. Leaf water potential at zero turgor ({Psi}L0) ranged from -1.76 MPa to -2.52 MPa before, and from -2.18 MPa to -2.59 MPa after water stress. Though {Psi}L0 of genotypes such as Cavalier, El Toro, and Emerald decreased after stress, {Psi}L0 of genotypes such as Korean Common and DALZ8515 did not change. Osmotic potential at full turgor ({Psi}{pi}100) of genotypes such as DALZ8501 and DALZ8506 was similar or increased after water stress but decreased for genotypes such as Crowne and Korean Common. The {Psi}L0 and {Psi}{pi}100 after stress were negatively correlated with recovery from stress and positively correlated with irrigation requirement. Zoysia genotypes with low relative water content at zero turgor (RWC0), bulk modulus of tissue elasticity ({epsilon}), and apoplastic water fraction (ß) demonstrated poor recovery from stress and required more supplemental irrigation. Cultivars such as Crowne, El Toro, and Palisades had the greatest recovery from stress, the least irrigation requirement, more negative {Psi}L0 and {Psi}{pi}100, and positive RWC0, {epsilon}, and ß. This study demonstrates that improvements in biophysical as well as morphological traits should contribute to development of water-conserving Zoysiagrass germplasm.

Abbreviations: {Psi}L0, water potential at zero turgor • {Psi}{pi}100, osmotic potential at full turgor • RWC0, relative water content at zero turgor • {epsilon}, bulk modulus of tissue elasticity • ß, apoplastic water fraction • TW:DW, turgid weight, dry weight ratio • {Delta}{Psi}p, change in turgor potential


   INTRODUCTION
TOP
 NOTES
 ABSTRACT
 INTRODUCTION
MATERIALS AND METHODS
 RESULTS AND DISCUSSION
 REFERENCES
 
WATER MANAGEMENT AND CONSERVATION continue to be major emphases of the turfgrass industry and turfgrass breeding programs. Water-use restrictions that limit irrigation of urban and suburban landscapes are commonplace. Therefore, development and use of cultivars with good drought resistance continues to be a major objective of turfgrass breeding programs.

Zoysiagrass, a C4 grass indigenous to the Orient, was introduced into the USA as recently as 1895 (Madison, 1971). Zoysiagrass has good winter-hardiness and high temperature tolerance (Beard, 1973) and is thus adapted to a wide environmental range. Zoysiagrass use in the USA has been limited because it usually requires propagation by sod, sod plugs, or sprigs. Slow establishment of many commercial varieties from sprigs or sod plugs escalates sod and sprig production costs, with subsequent high cost for establishment of most new turf sites. Aggressive breeding programs have developed new zoysiagrass germplasm with substantial improvements in establishment rates from sprigs and sod plugs. In addition, breeding efforts have recently focused on drought resistant genotypes.

Zoysiagrass is considered to be more drought resistant than C3 turfgrasses and intermediate to highly drought resistant compared with other C4 turfgrasses. The evapotranspiration rates of zoysiagrass cultivars and germplasm were similar under nonlimiting water availability (Green et al., 1991), but differed markedly in response to drought (White et al., 1993). Mean supplemental irrigation water required to prevent drought stress ranged from an average of 93 to 488 mm yr-1 among 21 zoysiagrasses during a 3-yr study.

Marcum et al. (1995) reported differences in rooting among 25 zoysiagrass genotypes with average maximum rooting depth from 151 to 381 mm under glasshouse conditions. Average maximum rooting depth, root mass, and root number at 300 mm were positively correlated with percentage green zoysiagrass ground cover in field study treatments that received no supplemental irrigation for 3 yr. Genetic differences in water requirement and drought resistance were evident among several Zoysia species. Although rooting characteristics were associated with field response to drought, published information concerning the physiological adaptation of zoysiagrasses to drought stress is lacking.

White et al. (1992) previously reported a strong relationship between turgor maintenance during soil drying and recovery of severely water-stressed tall fescue (Festuca arundinacea Schreb.) with low basal osmotic potential and osmotic adjustment contributing to turgor maintenance. Although evidence was provided to support a strong association between specific water relations characteristics and recovery from drought, limited published information is available concerning the association between water relations characteristics and water requirement of turfgrasses. Such information could provide a valuable basis for selection of germplasm with a low water requirement and superior drought resistance. The objectives of this work were to determine (i) water relations characteristics of 15 zoysiagrasses before and after water stress, (ii) survival and recovery from extreme water deficit stress, and (iii) the relationship to supplemental irrigation requirement of these same zoysiagrasses.


   MATERIALS AND METHODS
TOP
 NOTES
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
RESULTS AND DISCUSSION
 REFERENCES
 
Plant Culture
Fifteen zoysiagrass genotypes (Table 1) were selected on the basis of previous evaluation of drought stress response (White et al., 1993). Three sprigs of each genotype were propagated in 6.3-cm diameter by 6.3-cm-deep open bottom tubes filled with fritted clay. Tubes were supported in flats filled with fritted clay during the establishment period. After 16 wk, genotypes were transplanted to a 38-cm diameter by 45-cm-deep plastic tub containing 29 kg of fritted clay. Tubs had holes in the bottom for drainage. A total of four tubs (replications) were planted and subjected to water stress during July through February. A completely randomized experimental design was used. A complete set of the 15 genotypes were grown in the same container to (i) offset genotypic differences in water use that might affect rate of stress development (Thomas, 1987), and (ii) promote competitive soil water extraction among plants.


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  		Table 1. Plant recovery characteristics and annual supplemental irrigation requirement for zoysiagrass genotypes

 
Plants were grown in tubs for 12 wk, during which time plants were well watered and received weekly applications of 6 kg N, 2.6 kg P, and 5 kg K ha-1 from a soluble (20:8.74:17.8; N:P:K) fertilizer. Plants were preconditioned to water stress by withholding irrigation for 3 wk until soil moisture content was 0.38 g g-1. Field capacity of the fritted clay was about 0.64 g g-1. After rewatering to saturation, irrigation was withheld for 13 wk to create severe water deficit stress. At this point, plants had less than 5% green leaf tissue and soil moisture content was about 0.24 g g-1. Temperatures in the greenhouse during the 12 wk before stress and during the study ranged from 24 to 35EC. Daytime relative humidity ranged from 15 to 60%.

Water-Release Curves
Immediately before the preconditioning cycle (before water stress), three to six vertical stems were detached from each plant, recut under water, and hydrated in a vial of distilled water for 16 h in darkness at 5°C. Plants were sampled again similarly 35 d after initiating the stress cycle, at which time leaf lamina of all plants were severely rolled. A hydraulic press and sequential weighing technique (White et al., 1992) was used to generate leaf water potential ({Psi}L) and turgid weight and fresh weight values for the pressure–volume curve. Deviation from the previously reported procedure included the use of two to three most recently expanded leaves within replications rather than using a single leaf. Relative water content was calculated for each {Psi}L by:

where FW, DW, and TW are fresh weight, dry weight, and turgid weight. Data were plotted as 1/{Psi}L versus RWC (the pressure–volume curve) for each treatment combination within replications from samples before and after water stress was imposed. Osmotic potential at full turgor ({Delta}), {Psi}L0, RWC0, and the proportion of cell wall or bound water (ß) were derived from each curve (Hsiao et al., 1976). Bulk modulus of elasticity of lamina tissue ({epsilon}) was determined from the relationship between turgor potential ({Psi}ñ) and RWC assuming an approximate linear relation for {Psi}ñ > 0 (Wilson et al., 1979) by:

Turgid weight:dry weight ratios (TW:DW) were determined from initial weight after hydration and oven dry weight for each selection.

After 13 wk of stress, when plants had less than 5% green tissue, the herbage was cut at 2 cm above the growth medium surface, and dry weight of the harvest determined. Shoot dry weight after stress included the cumulative above-ground herbage mass produced during establishment and during water stress. Plants were then rewatered and fertilized. The number of new leaves produced, and growth, measured as dried, weighed herbage, were determined 4 to 6 wk later. Shoot recovery weight was thus the herbage produced during 4 to 6 wk after termination of water stress.

All data were subjected to analysis of variance. When a significant f-ratio occurred, Fisher's Protected Least Significant Difference was calculated for mean comparison.


   RESULTS AND DISCUSSION
TOP
 NOTES
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS AND DISCUSSION
REFERENCES
 
Annual supplemental irrigation requirement of the 15 zoysiagrass genotypes ranged from 93 to 488 mm (Table 1), as previously reported (White et al., 1993). The supplemental irrigation requirement for these zoysiagrasses was determined by the amount of irrigation required to prevent expression of drought stress symptoms such as wilting, leaf rolling, and loss of green pigmentation. As such, the annual supplemental irrigation requirement is an indicator of drought resistance among these zoysiagrasses (White et al., 1993). The same grasses were evaluated in the present study to determine the relationship between field response to drought and simulated drought under greenhouse conditions.

After-stress shoot weights differed among genotypes under greenhouse conditions (Table 1). In general, more robust plants, such as Crowne, Palisades, and El Toro, had greater after-stress shoot weights than more diminutive genotypes, such as Emerald and Diamond. After-stress shoot weights represented cumulative growth of genotypes during establishment and stress periods and as such may not have been a good indication of overall drought resistance. Shoot recovery weights were indicative of survival and growth following stress. Shoot recovery weight was greatest for Crowne and El Toro, and zero for DALZ8513, DALZ8501, and DALZ8515. New green leaves formed after stress may have also reflected growth rate, but new leaf formation was considered as an excellent indicator of survival of genotypes during severe water stress. El Toro produced the most new green leaves after termination of water stress followed by Crowne and Palisades. All other genotypes produced similar numbers of new green leaves. DALZ8501, DALZ8506, and DALZ8515 produced no new green leaves and had zero shoot recovery weight. These data indicated that the latter genotypes had poor drought resistance. Supplemental irrigation requirement was negatively correlated (P >= 0.01) with after-stress shoot weight, shoot recovery weight, and new green leaves (Table 2). These data indicated a strong relationship between field drought response and response to simulated drought under greenhouse conditions.


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  		Table 2. Correlation coefficients (r) among after stress shoot weight, recovery ratio, green leaf recovery, and supplemental irrigation requirement for 15 zoysiagrass genotypes

 
Significant water stress x genotype interaction effects occurred for all water relations characteristics except turgid TW:DW (Table 3). The {Psi}L0 ranged from -1.76 MPa for Diamond to -2.52 MPa for DALZ8515 before water stress (Table 3). Averaged across all cultivars, {Psi}L0 decreased after water stress; however, {Psi}L0 of genotypes such as Korean Common and DALZ8515 did not change during water stress. In contrast, {Psi}L0 of genotypes such as Cavalier, El Toro, and Emerald decreased substantially after water stress and {Psi}L0 ranged from -2.18 MPa for Diamond to -2.59 MPa for El Toro.


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  		Table 3. Water potential at zero turgor ({psi}L0), osmotic potential at full turgor ({Psi}{pi}100), relative water content at zero turgor (RWC0), bulk modulus of tissue elasticity ({epsilon}), apoplastic water fraction (ß), and turgid weight/dry weight ratio (TW:DW) determined before and after water stress, for 15 zoysiagrass genotypes

 
The {Psi}L0 for these zoysiagrasses were similar to those reported for tall fescue by White et al. (1992) but more negative than reported for tropical forage species reported by Wilson et al. (1980). Wilson et al. (1980) previously reported adjustments in {Psi}L0 during drought for green panic (Panicum maximum Jacq.), spear grass (Heteropogon contortus L.), and buffelgrass (Cenchrus ciliaris L.), although White et al. (1992) reported no adjustment in {Psi}L0 for tall fescue during drought. After 5 wk of water stress, average {Psi}L0 of all genotypes decreased by 0.17 MPa. The adjustment in {Psi}L0 that occurred is consistent with changes in {Psi}L0 reported by Wilson et al. (1980) but not by White et al. (1992) and Taylor (1998). The adjustment in {Psi}L0 observed in this study represents substantial genetic variation within each of these Zoysia species for physiological adaptation to soil water deficit.

The {Psi}L0 was negatively correlated with recovery weight after, but not before, water stress (Table 4). A strong positive relationship also existed between {Psi}L0 after water stress and irrigation requirement. The {Psi}L0 provides a quantitative indication of turgor loss point for comparison among genotypes or in the evaluation of specific physiological processes. Turgor maintenance during drought is considered of value for many plant species, contributing to more normal function or continuation of turgor mediated processes as plant available soil water becomes limited (Morgan, 1984). These data also established a strong relationship between turgor maintenance and supplemental irrigation water requirement. Genotypes with a more negative turgor loss point required significantly less supplemental irrigation. Marcum et al. (1995) previously reported a strong relationship between root morphology (number and maximum depth) and irrigation requirement of many of these same zoysiagrass genotypes. The current study is the first to demonstrate that irrigation water requirement is also under biophysical control in zoysiagrass.


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  		Table 4. Correlation coefficients (r) among water potential at zero turgor (QL0), osmotic potential at full turgor (QB100), relative water content at zero turgor (RWC0), bulk modulus of tissue elasticity (g), apoplastic water fraction ($), turgid weight/dry weight ratio (TW/DW) determined before and after water stress, and shoot weight, recovery weight, recovery ratio, and annual supplemental irrigation requirement for 15 zoysiagrass genotypes

 
The adjustment in {Psi}L0 was generally associated with {Psi}{pi}100, although RWC0, {epsilon}, ß, and TW:DW were also significantly correlated with {Psi}L0 both before and after water stress (Table 5). Osmotic potential and {Psi}L0 were positively correlated (r = 0.90, P >= 0.05) in tall fescue (White et al., 1992) after water stress. The range in {Psi}{pi}100 for zoysiagrass genotypes were similar to those reported for nonwater stressed Old World Bluestem (Bothriochloa spp.) by Coyne et al. (1982). After water stress, {Psi}{pi}100 for these zoysiagrasses was similar to water stressed tall fescue (White et al., 1992) and tropical forage grasses (Wilson et al., 1980), but much more negative than for creeping bentgrass (Agrostis palustris Huds.) (Taylor, 1998).


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  		Table 5. Correlation coefficients (r) among water potential at zero turgor (QL0), osmotic potential at full turgor (QB100), relative water content at zero turgor (RWC0), bulk modulus of tissue elasticity (g), apoplastic water fraction ($), turgid weight/dry weight ratio (TW/DW) determined before and after water stress, for 15 zoysiagrass genotypes

 
Osmotic potential at full turgor differed among genotypes before water stress and ranged from -1.18 MPa for Diamond to -1.76 MPa for Meyer and Palisades (Table 3). When averaged across all genotypes, {Psi}{pi}100 was similar before and after water stress. The significant water stress x genotype interaction effect indicates that some genotypes adjusted osmotically to a greater extent than other genotypes. Though {Psi}{pi}100 of genotypes such as DALZ8501, DALZ8506, and DALZ8515 was similar or even increased after water stress, {Psi}{pi}100 of genotypes such as Crowne, Korean Common, and Meyer decreased after water stress.

Some zoysia genotypes (DALZ8501 and DALZ8506) exhibited an increase in {Psi}{pi}100 after water stress, indicating a decrease in osmotic solutes. Decrease in osmotic solutes may have been due to (i) decrease in photosynthetic activity during stress because of stomatal sensitivity to water deficit or (ii) partitioning of osmotic solutes to other tissues. Poor shoot recovery weights and lack of new green leaf formation after water stress suggested a low energy state in DALZ8501 and DALZ8506, and indicated support for the former postulate.

Osmotic potential after water stress was negatively correlated with recovery weight and new green leaf formation, which indicated that more negative {Psi}{pi}100 improved survival and recovery (Table 4). A strong positive relationship existed between {Psi}{pi}100 and supplemental irrigation requirement. The {Psi}{pi}100 may prove to be a rapid and economical selection tool for drought resistant, water conserving Zoysia genotypes because osmotic potential is more easily and rapidly measured than the other water relations characteristics reported in this study.

Decreases in {Psi}{pi}100 during water stress may be accounted for by (i) changes in TW:DW, (ii) variation in the proportion of ß, and (iii) accumulation of solutes (Wilson et al., 1980). The TW:DW were similar among genotypes and was not affected by water stress (Table 3). Therefore, osmotic adjustment was not caused by changes in TW:DW. Differences in ß were observed among genotypes and water stress treatments. The ß changed an average of 0.10 g g-1 with a maximum of 0.18 g g-1 for DALZ8513. Wilson et al. (1980) reported an average increase in ß of about 0.10 g g-1 that accounted for 25, 31, and 21% of the osmotic adjustment that occurred in green panic, speargrass, and buffelgrass. They concluded that 70 to 80% of the osmotic adjustment occurred due to increases in osmotic solutes. Therefore, it was inferred that osmotic adjustment in Zoysia is primarily due to increases in osmotic solutes.

Mean zoysiagrass ß was negatively correlated with {Psi}{pi}100 both before and after water stress (Table 5). Although changes in ß probably affected changes in {Psi}{pi}100, ß probably contributed to a smaller percentage of the osmotic adjustment that occurred in these zoysiagrasses than did active solute accumulation (Wilson et al., 1980). Apoplastic water fraction was different among genotypes and water stress treatments (Table 3). Before water stress, ß ranged from 0.02 g g-1 for Diamond to 0.39 g g-1 for Korean common. After water stress, ß ranged from 0.16 g g-1 for DALZ8501 to 0.45 g g-1 for Palisades. The ß of genotypes changed disproportionately in response to water stress treatments as indicated by the significant water stress x genotype interaction effect. The ß for some genotypes, such as Korean Common and DALZ8501, did not change in response to water stress in contrast to other genotypes, such as Palisades and Diamond that had an increase in ß of about 0.15 units. Averaged over all genotypes, ß increased by 0.10 units due to water stress.

The {epsilon} was different for water stress treatments and among genotypes (Table 3). The significant water stress x genotype interaction effect reflects the substantial increase in {epsilon} (greater rigidity) by genotypes such as Meyer and Palisades in contrast to most of the Z. matrella that had similar or decreased {epsilon} (greater elasticity) after water stress compared to before water stress. Increased {epsilon} in zoysiagrass in response to drought is consistent with the response observed in several grass species (Wilson et al., 1980; White et al., 1992). The change in zoysiagrass {epsilon} in response to drought appeared to be species, as well as genotype, related. Among Z. japonica genotypes, all except one had increases in {epsilon} that ranged from 0.86 to 4.03 MPa in response to water stress; however, the Z. matrella genotypes exhibited divergent changes in {epsilon} decreasing as much as -1.68, and to an increase of 0.89 MPa. The TW:DW ratios were similar among genotypes, and were not associated with observed changes in {epsilon} (Tables 3 and 4); although, in general, absolute values of TW:DW declined after water stress compared with before stress. In some grasses, an increase in {epsilon} has been associated with a decrease in TW:DW ratios because of changes in cell-wall thickness or cell-wall constituents (Wilson et al., 1980).

Relative water content at zero turgor (RWC0) was similar before and after water stress (Table 3). The RWC0 before stress ranged from 0.66 g g-1 for Emerald to 0.82 g g-1 for El Toro and Korean Common. After water stress, RWC0 ranged from 0.67 g g-1 for Diamond and Emerald to 0.85 g g-1 for Palisades.

The RWC0, {epsilon}, and ß were negatively correlated with {Psi}L0 (Table 5) which indicated a strong influence of these characteristics on turgor maintenance. The RWC0, {epsilon}, and ß also demonstrated a strong negative association with supplemental irrigation requirement (Table 4). This study indicated that zoysiagrasses with lower {epsilon}, such as DALZ 8501, DALZ8506, and DALZ8515, also had lower RWC0 and ß (Tables 3 and 5). Zoysia genotypes with low RWC0, {epsilon}, and ß, in general, had a high annual supplemental irrigation requirement and demonstrated poor drought resistance as indicated by less shoot recovery weight and new green leaves formed after stress. Therefore, Zoysia genotypes with low RWC0, {epsilon}, and ß may not contribute to improvements in drought resistance. In general, the greatest recovery from stress and the least annual supplemental irrigation requirement was observed in grasses such as Crowne, El Toro, and Palisades that had more negative {Psi}L0, and {Psi}{pi}100, and more positive RWC0, {epsilon}, and ß.

Though this study did not definitively delineate the degree of genetic control of water relations characteristics, annual supplemental irrigation requirement, and drought resistance, strong evidence is presented that a great degree of genetic variation exists for these traits among these Zoysia genotypes. The data presented demonstrated that supplemental irrigation requirement was under biophysical control in Zoysia and that improvements in biophysical as well as morphological traits may contribute to the development of more drought resistant, water-conserving Zoysia germplasm.


   NOTES
TOP
 NOTES
ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS AND DISCUSSION
 REFERENCES
 
Supported in part by a grant from the U. S. Golf Association - Green Section.

Received for publication October 25, 1999.


   REFERENCES
TOP
 NOTES
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS AND DISCUSSION
 REFERENCES
 




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