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Dep. of Horticulture, Forestry, and Recreation Resources, Kansas State Univ., Manhattan, KS 66506-5506 USA
bhuang{at}oz.oznet.ksu.edu
| ABSTRACT |
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Abbreviations: Fv/Fm, photochemical efficiency LSD, least significance difference Pn, canopy photosynthetic rate
| INTRODUCTION |
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The lower optimum temperature for root growth than for shoot growth of cool-season grasses indicates that roots may be more sensitive to high temperatures. (Beard, 1973). Root growth of Kentucky bluegrass (Poa pratensis L.) was inhibited as soil temperature increased to 25°C (Aldous and Kaufmann, 1979). Root growth and initiation in creeping bentgrass stopped when soil temperature was above 25°C (Lucas, 1995; Beard and Sifers, 1997; Huang et al., 1998). Soil temperature strongly influenced shoot growth of winter wheat (Triticum aestivum L.) by regulation of cytokinin production in roots (Kuroyanagi and Paulsen, 1988). Direct injury to roots by high soil temperatures could well be the initial factor in high temperature responses of plants.
Reducing soil temperature by any means may alleviate or prevent the summer bentgrass decline problem. Reducing root-zone temperature has been shown to increase root growth, export of cytokinin from roots, leaf photosynthesis, chlorophyll content, protein synthesis, and shoot growth in several plant species (Skene and Kerridge, 1967; Feierabend and Mikus, 1977; Aldous and Kaufmann, 1979; Martin et al., 1985; Kuroyanagi and Paulsen, 1988; Graves et al., 1991; Clarck and Reinhard, 1991; Udomprasert et al., 1995; Ziska, 1998). However, whether shoot and root growth of creeping bentgrass can be improved by reducing root-zone temperature when shoots are exposed to supraoptimal air temperatures, has not been studied. Furthermore, the relative effects of air compared with soil temperatures on creeping bentgrass are not well understood.
The objectives of this study were (i) to compare the influences of air versus soil temperature on turf quality, physiological activities, and root growth and (ii) to investigate whether growth and turf quality of creeping bentgrass could be improved by reducing soil temperature under high ambient temperature conditions.
| Materials and methods |
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Shoots and roots were exposed to four air/soil temperature regimes: 20/20, 20/35, 35/20, and 35/35°C for 56 days. These test temperatures were chosen because 20°C is within the optimum temperature range for the growth of cool-season grasses, and 35°C commonly occurs in the transitional and warm climatic regions during mid-summer. Shoots were maintained at the ambient temperature (20 or 35°C) of the growth chambers, which was regulated by a temperature controller. Soil temperature was controlled by maintaining the entire root zone (40-cm-long soil column in a polyethylene bag) in water baths with predetermined temperatures (20 or 35°C). The low temperature was controlled with circulating cool water (1820°C), and the high temperature with an immersion circulating heater (IC-2100, Fisher Scientific Inc., Pittsburgh, PA). Water was circulated continuously to maintain constant and uniform temperatures. Water levels were maintained at the top edges of both water baths during the experimental period.
Air temperatures at various distances from the canopy and soil temperatures at different depths from the soil surface were measured biweekly with thermocouples connected to a thermometer (Fig. 1) . In the 20/20°C treatment, both air and soil were maintained around 20°C; in the 20/35°C treatment, air temperature ranged from 23 to 25°C, and root-zone temperature was approximately 35°C; in the 35/20°C treatment, air temperature ranged from 32 to 35°C, while root-zone temperature ranged from 18 to 21°C; and in the 35/35°C treatment, air and root-zone temperatures were maintained at 33 to 36°C.
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Turf quality was visually rated weekly after treatments were initiated, and was based on color, density, and uniformity on a 0 (worst, all plant were dead and brown) to 9 scale (best, all plants were healthy and green), with scales above 6 being acceptable quality. Leaf photochemical efficiency was expressed as leaf chlorophyll fluorescence (Fv/Fm). The Fv/Fm ratio of leaves was determined with a chlorophyll fluorescence meter (Dynamax, Houston, TX) by pressing the light source chamber to leaves that were adapted in dark for 20 min.
Diurnal gas exchange rate was measured every 2 h from 0900 to 1300 h on 28 d of treatment with a LI-6400 portable gas exchange system (LI-COR Inc., Lincoln, NE). Turf canopy was enclosed in a transparent plexiglass chamber fitted to the CO2 analyzer in LI-6400. Canopy net photosynthetic rate (Pn) was expressed as CO2 uptake per unit turf canopy area (m-2). Changes in canopy photosynthetic rate (Pn) with duration of treatment was also determined by measuring Pn from 10:00 to 14:00 h weekly using the LI-6400 gas exchange system.
The number of roots at different soil depths was determined nondestructively with an imaging system. Roots visible on the side wall of the transparent polyethylene bags were recorded at depths of 0 to 2, 5 to 7, 15 to 17, 25 to 27, and 35 to 25 cm from the soil surface using a video-camera recorder (Sony, CCD-TRV70, Japan). The number of roots within a 2- by 3-cm area at each soil depth was determined from the root images.
Plants from four tubes in each treatment were harvested biweekly to determine root fresh weight. Fresh weight was determined after roots were washed and blotted dry with paper towels.
| Results |
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Canopy Photosynthetic Rate
Canopy photosynthetic rate (Pn) for grasses grown at 20/20°C and 35/20°C remained relatively constant during the experimental period (Fig. 3)
. However, Pn declined to below the control level within 8 d after roots (20/35°C) and 1 d after both shoots and roots (35/35°C) were exposed to high temperature and continued to decline with treatment duration. The decline in Pn was more severe at 35/35°C than at 20/35°C. The Pn at 35/20°C was higher than that at 20/35°C and 35/35°C, but was not different from that at 20/20°C during the experiment.
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Root Fresh Weight
Root fresh weight declined with duration of treatment at 20/35, 35/20, and 35/35°C. This decline was more dramatic at 20/35 and 35/35°C than at 35/20°C (Fig. 6)
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Root Number
The number of roots decreased with soil depths in each treatment (Fig. 7)
. The numbers of roots remained constant during the experimental period in all soil depths at 20/20°C, in soil depths below 5 cm at 35/20°C, and at depths below 35 cm at 20/35°C (Fig. 7). Significant declines in the root number with treatment duration occurred in the 0- to 2-cm depth at 20/35, 35/20, and 35/35°C; in 5- to 7-, 15- to 17-, and 25- to 27-cm depths at 20/35 and 35/35°C; and in all soil depths at 35/35°C. However, the magnitude of the decline in root number was greater in the surface 7 cm soil than in the deeper soil layers. The decrease in the number of roots in the 0- to 2-cm depth at 35/35°C was more dramatic than decrease in all other treatments. The declines in the number of roots in soil depths below 5 cm were greater at 20/35 and 35/35°C than at 35/20 and 20/20°C.
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| Discussions |
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The adverse effects of high soil temperature on shoot growth and physiological activities probably are due to direct inhibition of root growth and activity and, therefore, limitation of water and nutrient supplies to the shoot (Kramer, 1983), and disruption of cytokinins synthesis in roots (Itai et al., 1973; Al-Khatib and Paulsen, 1984; Kuroyanagi and Paulsen, 1988; Smart et al., 1991). High soil temperature also promoted leaf senescence by increasing transport of root abscisic acid (ABA) to the shoots (Udomprasert et al., 1995).
High soil temperature or high air and soil temperatures reduced fresh weight and number of roots in different soil depths and accelerated the decline in root weight and number. The decline in the number of roots was more pronounced in the surface 7 cm of soil where most roots were found. Rapid decline in the number of roots under high soil temperatures suggested that root death rate exceeded the rate of new root production, especially in the surface soil layers. Beard and Daniel (1965) reported that no new root growth of creeping bentgrass occurred in the summer, except during two low temperature periods. Aldous and Kaufmann (1979) reported that death of the roots under heat stress initiated the decline of shoot growth in Kentucky bluegrass, which might result from a high temperature-labile process in the roots.
Limited new root production and accelerated root death under high root temperatures may interrupt synthesis and transport of root-produced hormones. Cytokinin is a shoot growth regulator that is synthesized mainly in roots (Skene and Kerridge, 1967). Its synthesis in roots and translocation from roots to shoots have been reported to be inhibited by high soil temperatures. Itai et al. (1973) and Skene and Kerridge (1967) reported adverse effects of high soil temperature on the exudation of cytokinins from detached roots. Heat stress damages to shoot chloroplast activities and ultrastructure are reversed by cytokinin (Caers et al., 1985). Udomprasert et al. (1995) reported that high soil temperature decreased endogenous ABA levels in roots of beans (Phaseolus vulgaris L.), but increased ABA in leaves, possibly by higher transport of root ABA to the shoots, which can cause leaf senescence.
Reducing soil temperature while shoots were exposed to high air temperature increased photosynthetic rate, photochemical efficiency, turf quality, and root growth, relative to plants grown at high soil temperatures. The effect on root growth was indicated by the slow decline or unchanged root fresh weight and the number of roots in different soil depths during high temperature stress to shoots. The increased growth and activity from reducing root temperature could improve the ability of roots to supply water (Kramer, 1983), nutrients (Engels, 1993), and hormones (Skene and Kerridge, 1967) for shoot growth.
Enhancement of shoot growth and physiological activities by reducing soil temperature have been reported in several other species (Aldous and Kaufmann, 1979; Martin et al., 1985; Kuroyanagi and Paulsen, 1988; Ruter and Ingram, 1990; Graves et al., 1991; Udomprasert et al., 1995; Ziska, 1998). Kuroyanagi and Paulsen (1988) reported that cooling roots of winter wheat to 25°C when shoots were exposed to 35°C was nearly as beneficial as growing both roots and shoots at 25°C. Shoot growth rate, specific leaf weight, leaf carbon exchange rate, and stomatal conductance in several plants were enhanced by low soil temperature regardless of the air temperature (Martin et al., 1985; Ziska, 1998). Survival of Kentucky bluegrass was enhanced by controlling soil temperature at 22°C (Aldous and Kaufmann, 1979). Low soil temperature also improved shoot extension in two woody species (Martin et al., 1985).
In summary, this study, along with others, demonstrated that roots had more influence than shoots on whole-plant responses to heat stress. Lowering soil temperature even when air temperature was supraoptimal enhanced growth and physiological activities of shoots and roots. Therefore, any means that reduce soil temperature during hot summers could improve turf quality of creeping bentgrass. The mechanisms for the mediation of high-temperature injury by roots in cool-season turfgrasses deserve investigation.
| ACKNOWLEDGMENTS |
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Received for publication September 23, 1999.
| REFERENCES |
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