(0038-075X,/92/1544.0290803.00/0 Sou. Science Copyright © 1992 by Williams & Wilkins October 1992 Vol. 154, No. 4 Printed in U.S.A. SOIL TEMPERATURE AND ROOT GROWTH! TC, Soil temperature affects both the rate and thoroughness with which a plant root system permeates soil. Root system expan- sion is a function of two temperature- dependent processes, growth and develop- ment. Growth processes, like cell elonga- tion, increase root length and diameter. Development controls duration of growth and initiation of new roots and reproduc- tive organs. Interpreting root temperature responses requires an understanding of how development and growth interact. Soil temperature affects growth of root system components, initiation and branching, ori- entation and direction of growth, and root turnover. Genotypic differences in root re- sponse to soil temperature exist between and within plant species. In natural soil profiles, root system expansion is affected by seasonal patterns of soil temperature. As soil warming advances downward, pro- gressively deeper soil layers become suit- able for root growth. In temperate regions, soil temperature often limits the rate of rooting-depth increase and the maximum depth attainable. A simple temperature- based model to predict rooting depth with time indicates that rooting depth may fol- low the downward progression of a partic- ular isotherm, which has sometimes been observed in the field. ‘Temperature is an important environmental regulator of plant growth. In temperate regions, the seasonal climate pattern determines when temperatures are suitable for growing warm- season crops. Aboveground, this seasonal pat- tern is predominantly temporal; belowground, temperature limitations to growth have both, temporal and spatial (depth) boundaries (Fig. ‘Joint contribution from the National Soil Tilth Lab., USDA-ARS, and the Dept. of Soil Science, Uni- versity of Wisconsin-Madison, * National Soil Tilth Laboratory, 2150 Pammel Dr., ‘Ames, IA 50011 * Department of Soil Science, University of Wiscon- sin-Madison, 1525 Observatory Dr., Madison, WI, 53706-1299, Received June 22, 1992; accepted June 24, 1992 KASPAR? ann W. L, BLAND® 1). As soil warming advances downward during the growing season, progressively deeper soil layers become suitable for root growth. The depth of soil thermally suitable for appreciable root growth is a function of both soil tempera- ture regime and root temperature response. ‘Thus, soil temperature often limits both root system expansion and proliferation, particularly during the early growing season. Our understanding of how roots interact with temporal and spatial patterns of soil tempera- ture is still limited. For 20 years, reviewers have written of the need to study root system expan- sion and development in thermal environments typical of cropping systems (van Bavel 1972; Bowen 1991). Factors impeding this research include the difficulty of studying plant roots in soil and the challenges of recreating soil thermal regimes under controlled conditions. This re- view first addresses how temperature response of roots can be altered by interactions between growth and development processes and by ex- perimental and environmental factors. Next, temperature effects on growth of root system components, initiation and branching, orienta- tion and direction of growth, and root turnover are discussed. Lastly, the hypothesis that rate of rooting depth increase is determined by the influence of soil temperature regimes on the elongation of the deepest roots is considered. Interactions of temperature with aeration and water and nutrient uptake are not discussed. The terms soil temperature and root tempera- ture are used interchangeably. INTERACTIONS BETWEEN GROWTH AND DEVELOPMENT. Root system expansion is a function of two temperature-dependent processes, growth and development. Growth processes, like cell elon- gation, increase root length and diameter. De- velopment controls duration of growth processes and initiation of new roots and reproductive organs, Increases in total root length are the result of both root elongation and initiation of new roots. To interpret the effect of temperature on root systems, the interaction of growth and development must be considered. 290SOIL TEMPERATURE AND ROOT GROWTH Temperature (°C) T 0.0 Depth (m) 2.0) ‘8000, r 120 140 160 180 200 220 Day of year 240 Fic, 1. The seasonal warming of air temperatures (1a; 5-year average of daily means) determines when temperatures are favorable for plant growth, (eg, plants requiring air temperatures above 20°C begin growth after day 100). Soil temperatures favorable for root growth vary with time and depth (1b; modeled thermal regime). For example, root growth requiting temperatures greater than 20°C are possible at times and depths to the right and sbove the 20°C isotherm. Developmental processes generally control growth duration of cells (Burstrom 1956; Beau- champ and Lathwell 1966), organs, and whole plants (Beauchamp and Lathwell 1967). For ex- ample, Burstrom (1956) found that final root cell length of wheat (Triticum aestivum L.) de- creased with increasing temperature, even though elongation rate increased with tempera- ture. Cells were shorter because development rate increased with increasing temperature, re sulting in a shorter growth duration. Growth is often reduced by a lack of water (Brouwer and van Vliet 1960; Markhart et al. 1979; Kramer 1983), nutrients (Brouwer and van Vliet 1960), or photosynthate (Rufty et al. 1981; Szaniawski and Kielkiewiez, 1982; Gregory 1986; Rice and Bastin 1986). In contrast, plant development rate usually is less sensitive to these factors (Monteith 1981; Thorne and Wood, 1987). When root growth is limited by a factor other than temperature, development may proceed at a rate limited only by tempera- ture, resulting in a smaller than normal root system. This difference between root growth rate and development rate can be especially critical just before the onset of seed filling, 291 INTERACTION WITH EXPERIMENTAL AND ENVIRONMENTAL FACTORS Interpretation of root temperature response is often complicated by interactions with experi- mental conditions and environmental factors. In many early studies, soil temperature was not controlled independently of air temperature (Richards et al. 1952; Nielsen and Humphries 1966), confounding their effects. Root response to temperature also changes with duration of the temperature treatment (Brouwer and van Viiet 1960; Brouwer 1962; Brouwer 1964) or with pretreatment temperature regimes (Brouwer 1964). When environmental or experimental factors other than temperature affect root growth, re- sponse of roots to soil temperature may be al- tered. In some cases, alteration of the tempera- ture response cannot be explained by interac- tions between growth and developmental processes. Some factors that alter temperature response include P concentration (Nielsen et al. 1960b; Case et al. 1964; MacKay and Barber 1984), rooting media (Abbas Al-Ani and Hay 1983), shoot temperature or light levels (McAdam and Hayes 1981; Rufty et al. 1981; Loffroy et al. 1983; MacKay and Barber 1984), soil water (Mack and Finn 1970), and soil strength (Pearson et al. 1970). Figure 2 shows how phosphorus fertilizer enhanced the root temperature response of oats (Avena sativa L.). As a result, comparison of data and conclusions from different experiments must be considered carefully, Root Dry Weight (g/plant) ‘Temperature (°C) Fic, 2. Root temperature response of oat root dry ‘weight per plant at heading for three fertilizer treat- ‘ments, (Drawn from the data of Nielsen et al. 1960b.)292 ROOT DRY WEIGHT Dry weight is the most commonly measured root parameter (Cooper 1973). Since 1973, tem perature effects on root dry weight have been reported for a range of species including blue grama [Bouteloua gracilis (Willd. ex H.B.K.) Lag ex Griffiths; Wilson 1981], soybean [Glycine ‘max (L.) Mert; Matthews and Hayes 1982), perennial ryegrass (Lolium perenne L. Clarkson, et al, 1986), and sorghum (Sorghum vulgare; Rao et al. 1989). In general, root dry weight follows a typical temperature response curve Fig. 3) Plant response to soil temperature differs among genera (Cooper 1973). Brouwer (1962) examined the seedling root growth of eight erop species (Table 1). All eight species had optimum, root growth at about 25°C, whereas other studies, cited by Cooper (1973) showed species differ- ences for optimum temperature. The eight spe- cies in Brouwer's study, however, differed in their response to temperatures above and below optimum {Table 1). Plant species within the same genus (Medicago; Nielsen et al. 1960a) and genotypes within species [tobacco (Nicotiana ta- bacum L.)—Johnson and Hartman 1919; bro: megrass (Bromus inermis Leyss.)—Dibbern 1947; perennial ryegrass—Kleinendorst and Brouwer 1965; timothy (Phleum pratense L.)— ‘Mack and Finn 1970; maize (Zea mays L.)—Cal and Obendorf 1972; Frossard 1985] also re- sponded differently to soil temperature. Root Weight (% of 25°C) i L 5 10 18 20 25 30 35 40 Root Temperature (°C) Fic. 3, Root temperature response of maize root ry weight at 24 days after germination. (Redrawn from Brouwer 1962.) KASPAR AND BLAND TABLE 1 Root temperature range for root growth rates greater than or equal to 50% of the maximum Species range Tow Flax (Linum usitatissimum L) 10 Peas (Pisum sativum L.) 9 Kidney bean (Phaseolus vulgaris L.) 12 Maize (Zea mays L.) uw Strawberry (Frogaria sp.) 5 Broad bean ( Vieia faba) 2 Rape (Brassica napus L.) 16 Oat (Avena sativa L.) 9 Data are from Brouwer (1962); source Klepper (1987), reproduced by permission of Cambridge Uni: versity Press. ROOT LENGTH In the past, root length was difficult to mea. sure and was infrequently reported. However, root length, surface area, and extension of roots into unexploited soil greatly influence water and nutrient uptake. Using a mechanistic P uptake model, Barber et al. (1989) showed that root: elongation rate was the most important model parameter for P uptake from soil Elongation of individual roots responds to temperature in much the same way as does root dry weight (Fig. 4). Recent studies of root length, response to temperature include Atkin et al 1973 and Blacklow 1972—maize; Cumbus and Nye 1982—rape (Brassica napus L.); Abbas Al- Ani and Hay 1983—wheat, barley (Hordeum sativum L.), rye (Secale cereale L.) and oats; Stone and Taylor 1983—soybean; Macduff et al. 1986—barley and rape; and Miyasaka and Grunes 1990—wheat. Potato (Solanum tubero- sur) and soybean genotypes also differed within species in their root elongation temperature re- sponse (Stone and Taylor 1983; Sattelmacher et al. 1990). Root length can be more sensitive to temper- atures below the optimum than root dry weight. Loffroy et al. (1983) found that cotton (Gos- sypium hirsutum L.) taproot length at a root temperature of 17°C was less than 40% of its length at 27°C, whereas root dry weight at 17°C was 68% of that at 27°C, Macduff et al. (1986) observed that root dry weight of rape was af- fected less by decreasing temperature below 7°C.