Effects of temperature on parameters of root growth relevant to nutrient uptake: Measurements on oilseed rape and barley grown in flowing nutrient solution

Summary Effects of root temperature on the growth and morphology of roots were measured in oilseed rape (Brassica napus L.) and barley (Hordeum vulgare L.). Plants were grown in flowing solution culture and acclimatized over several weeks to a root temperature of 5°C prior to treatment at a range of root temperatures between 3 and 25°C, with common shoot temperature. Root temperature affected root extension, mean radius, root surface area, numbers and lengths of root hairs. Total root length of rape plants increased with temperature over the range 3–9°C, but was constant at higher temperatures. Root length of barley increased with temperature in the range 3–25°C, by a factor of 27 after 20 days. Root radii had a lognormal distribution and their means decreased with increasing temperature from 0.14 mm at 3°C to 0.08 mm at 25°C. The density of root hairs on the root surface increased by a factor of 4 in rape between 3 and 25°C, but in barley the highest density was at 9°C. The contribution of root hairs to total root surface area was relatively greater in rape than in barley. The changes in root system morphology may be interpreted as adaptive responses to temperature stress on nutrient uptake, providing greater surface area for absorption per unit root weight or length.     

Phytophthora cryptogea root rot of tomato in rockwool nutrient culture:

The effect of root temperature on growth and yield of rockwool-grown tomato plants infected with Phytophthora cryptogea was investigated. Measurements of shoot and root growth were taken at high (25^oC) and low (15^oC) root temperatures during the generative phase of growth. The growth of roots of healthy and P. cryptogea-infected tomato plants in rockwool blocks was higher in plants grown with roots at 25^oC than at 15^oC after 60 days and a similar effect was found in slabs after 98 days. Under sub-optimal conditions for growth the disease became severe when root temperatures were low. Growth of roots was greatest when roots were maintained at a high temperature in combination with an ambient air temperature of c. 15^oC and the response was greater in cv. Counter than cvs Calypso and Marathon. Water-soluble carbohydrates of roots were higher in those produced in blocks than slabs and were reduced by infection compared to healthy plants with roots at 15^oC and 25^oC. Reduced transpiration rates were found 17 days after inoculation in symptomless plants grown at a root temperature of 25^oC. Infection, regardless of the temperature of the roots or cultivar, led to reduced stem growth. The plants grown at 25^oC were taller than those with a root temperature of 15^oC. After 9 wk of harvest, the cumulative fruit yields in infected cvs Counter and Calypso grown at 25^oC were comparable to that in healthy plants grown at either temperature and cumulative fruit numbers followed a similar pattern. High root temperatures led to delayed fruit ripening between weeks 3–10 and a larger number of unripe fruit. The weight of unripe fruit from infected plants grown at 25^oC at the terminal harvest was higher than from healthy plants with roots maintained at 15^oC.     

Arbuscular mycorrhiza colonization and development at suboptimal root zone temperature

Temperature has a strong influence on the activity of living organisms. This study, involving two indoor experiments, evaluated the effects of root zone temperature (10, 15 and 23°C) on the formation and development of arbuscular mycorrhizae (AM). In the first trial, greenhouse-grown sorghum [Sorghum bicolor (L.) Moench] was either colonized by Glomus intraradices Schenck & Smith or left non-mycorrhizal. Root length, root and shoot weight and root colonization were measured after 5, 10 and 15 weeks of plant growth. Although suboptimal root zone temperatures reduced growth in both mycorrhizal and non-mycorrhizal plants, mycorrhizal plants were larger than non-mycorrhizal plants after 15 weeks at 15 and 23°C. At suboptimal root zone temperatures, mycorrhizal inoculation sometimes slightly reduced root development. AM colonization was more affected than root growth at suboptimal root zone temperatures. Colonization was markedly reduced at 15°C compared with 23°C, and almost completely inhibited at 10°C. The second experiment was conducted in vitro using transformed carrot (Daucus carota L.) roots supporting G. intraradices. Mycelium length and spore number were measured weekly for 15 weeks. Spore metabolic activity (iodonitrotetrazolium reduction), root length and percentage root colonization were measured after 15 weeks. G. intraradices sporulation was reduced at temperatures below 23°C, while spore metabolic activity was significantly reduced only at 10°C. Root length and in particular percentage colonization were decreased at suboptimal temperatures. A negative interaction between AM hyphal growth and root growth resulting in reduced probability of contact at suboptimal root zone temperatures is proposed to explain the greater reduction observed in root colonization than in root and hyphal growth.     

Nitrogen utilization by Typha latifolia L. as affected by temperature and rate of nitrogen application

Greenhouse and growth chamber studies were conducted to evaluate growth and N utilization by Typha latifolia L. in flooded organic soil under varying temperatures and rates of N additions. Elevation of temperature from 10 to 25°C increased shoot biomass yields by 275%. Root biomass yields were lowest at 10°C and increased linearly as a function of temperature. Shoot/root ratios were low (0.72–0.82) at lower temperatures (10–15°C) and ratios increased by about three times at higher temperatures (20–30°C). Biomass yields were increased by addition of N fertilizers, while the shoot/root ratios were directly related to plant-available N present in the soil.Fertilizer ¹⁵N uptake (expressed as % of applied N) by the whole plant was 5.3% at 10°C, 37.5% at 20°C and at 30°C decreased to 20.8%. Fertilizer N accumulation in shoots was 2.1–29.8% of applied N, while roots accumulated 3.2–7.7%. Under greenhouse conditions, N uptake by T. latifolia was found to increase with increased rate of N application. Fertilizer N uptake by both shoots and roots was in the range of 61–77%. Plants cultured in growth chambers were affected by low light conditions resulting in poor growth and low fertilizer ¹⁵N uptake, as compared to plants grown under greenhouse conditions. Added fertilizer N was the major source of N during the early part of the growing season, while soil organic N was the major and perhaps the sole source of N during the latter part of the growing season.     

Maize root response to N in the greenhouse and field

Summary Root growth and morphology were compared between seven week old maize plants grown in the greenhouse and in the field. The plants were similar in shoot dry weight and the partioning of N and dry matter to roots were similar except for the field grown plants in 1983. Field grown plants had greater root mass per length and greater calculated diameter than greenhouse plants. Nitrogen fertilization decreased N and dry matter partitioning to the root system in all three environments.     

Influence of temperature on root development and phosphate influx in winter wheat grown at different P levels

Root development was studied in winter wheat ( Triticum aestivum L. cv Starke II) grown at 5,10, 15 and 20°C in nutrient solutions with phosphate concentrations of 10, 100 or 1000 μM . The plants were grown for 38 days (5 and 10°C), 19 days (15°C) or 14 days (20°C). At the end of the cultivation period the phosphate influx in the roots was determined with ³²P-phosphate. Root development (lateral and seminal roof length and number) was monitored throughout the cultivation period on the same individuals by repeated (approximately every second day) photocopying of the roots for measurements with digitizer and appropriate software. The 5°C treatment yielded no laterals, and the seminals were only slightly affected by the different phosphate treatments. The 10 μM phosphate treatment gave high root:shoot dry weight ratio, high average lateral root length and high specific root length [m root (g root fresh weight)^-1]. The 1000 μM phosphate treatment yielded the highest number of laterals per m seminal root, and usually also the highest absolute numbers. Phosphate influx decreased with increased P status of the roots. It is argued that phosphate influx is dependent on factors such as P status, root geometry and relative root extension rate.     

An assessment of the role of ethylene in mediating lettuce (Lactuca sativa) root growth at high temperatures

Growth of temperate lettuce (Lactuca sativa) plants aeroponically in tropical greenhouses under ambient root-zone temperatures (A-RZTs) exposes roots to temperatures of up to 40 degrees C during the middle of the day, and severely limits root and shoot growth. The role of ethylene in inhibiting growth was investigated with just-germinated (24-h-old) seedlings in vitro, and 10-d-old plants grown aeroponically. Compared with seedlings maintained at 20 degrees C, root elongation in vitro was inhibited by 39% and root diameter increased by 25% under a temperature regime (38 degrees C/24 degrees C for 7 h/17 h) that simulated A-RZT in the greenhouse. The effects on root elongation were partially alleviated by supplying the ethylene biosynthesis inhibitors aminooxyacetic acid (100-500 microM) or aminoisobutyric acid (5-100 microM) to the seedlings. Application of the ethylene precursor 1-aminocyclopropane-1-carboxylic acid to seedlings grown at 20 degrees C mimicked the high temperature effects on root elongation (1 microM) and root diameter (1 mM). Compared with plants grown at a constant 20 degrees C root-zone temperature, A-RZT plants showed decreased stomatal conductance, leaf relative water content, photosynthetic CO(2) assimilation, shoot and root biomass, total root length, the number of root tips, and root surface area, but increased average root diameter. Addition of 10 microM ACC to the nutrient solution of plants grown at a constant 20 degrees C root-zone temperature mimicked the effects of A-RZT on these parameters but did not influence relative water content. Addition of 30 microM aminoisobutyric acid or 100 microM aminooxyacetic acid to the nutrient solution of A-RZT plants increased stomatal conductance and relative water content and decreased average root diameter, but had no effect on other root parameters or root and shoot biomass or photosynthetic CO(2) assimilation. Although ethylene is important in regulating root morphology and elongation at A-RZT, the failure of ethylene biosynthesis inhibitors to influence shoot carbon gain limits their use in ameliorating the growth inhibition induced by A-RZT.     

Mediation of high-temperature injury by roots and shoots during reproductive growth of wheat

Abstract Previous studies suggest that high temperature stress on wheat (Triticum aestivum L.) involves root processes and acceleration of monocarpic senescence. Physiological changes in wheat roots and shoots were investigated to elucidate their relationship to injury from elevated temperatures after anthesis. Plants were grown under uniform conditions until 10 d after anthesis, when shoot/root regimes of 25°C/25°C, 25°C/35°C, 35°C/25°C and 35°C/35°C were imposed. Growth and senescence of shoots and grain were influenced more by root temperatures than by shoot temperatures. High root temperatures increased activities of protease and RNasc enzymes, and loss of chlorophyll, protein and RNA from shoots, whereas low root temperatures had opposite effects. High root temperatures appeared to induce shoot senescence directly. High shoot temperatures probably disrupted root processes, including export of cytokinins, and induced high leaf protease activity, senescence and cessation of grain development. The authors concluded that responses of wheat to high temperatures, whether of roots or shoots, are manifested as acceleration of senescence and may be mediated by roots during grain development.     

The effect of photoperiod and temperature on growth and frost resistance of winter rye root systems

Root growth, development and frost resistance were examined in winter rye ( Secale cereale L. cv. Puma) plants grown under 6 combinations of temperature and photoperiod (20/16°C or 5/3°C, day/night; 8, 16- or 24-h days). Overall root system growth is influenced by the interaction of temperature and photoperiod. Maximum shoot growth occurs at a 24-h photoperiod in 20°C plants and at a 16-h photoperiod in 5°C plants, and is correlated in both treatments with a high root:shoot ratio. Frost resistance of rye roots is affected by short photoperiods in 2 ways. First, short photoperiod and low temperature delay production of new adventitious roots so that newly developing roots are not exposed to freezing temperatures. Second, short photoperiod alone can induce several degrees of frost tolerance in existing roots during the lag phase of growth. Low temperature alone does not decrease the rate of dry weight accumulation in rye root systems, but cold temperature does retard developmental processes within the roots. Rye roots grown at 5°C develop first order lateral roots, differentiate metaxylem vessels and suberize endodermal cell walls more slowly than roots grown at 20°C.     

The influence of waterlogging at different temperatures on penetration depth and porosity of roots and on stomatal diffusive resistance of pea and maize seedlings

The 8 days old seedlings of pea (cv. Ilowiecki) and maize (cv. Alma F1) were subjected to differentiated aeration conditions (control — with pore water tension about 15 kPa and flooded treatment) for 12 days at three soil temperatures (7, 15 and 25 °C). The shoots were grown at 25 °C while the soil temperature was differentiated by keeping the cylinders with the soil in thermostated water bath of the appropriate temperature. Lowering the root temperature with respect to the shoot temperature caused under control (oxic) conditions a decrease of the root penetration depth, their mass and porosity as well as a decrease of shoot height, their mass and chlorophyll content; the changes being more pronounced in maize as compared to the pea plants. Flooding the soil diminished the effect of temperature on the investigated parameters; the temperature effect remaining significant only in the case of shoot biomass and root porosity of pea plants. Root porosity of pea plants ranged from 2 to 4 % and that of maize plants — from 4 to 6 % of the root volume. Flooding the soil caused an increase in the root porosity of the pea plants in the entire temperature range and in maize roots at lower temperatures by about 1 % of the root volume. Flooding the soil caused a decrease of root mass and penetration depth as well as a decrease of plant height, biomass and leaf chlorophyll content.     

Root contraction helps protect the "living rock" cactus Ariocarpus fissuratus from lethal high temperatures when growing in rocky soil

? Premise of the study: We investigated how the "living rock" cactus Ariocarpus fissuratus, like other low-growing desert plants, can endure potentially lethal high temperatures at the soil surface. Specifically, we examined how shoot descent by root contraction in the presence or absence of soil rocks influences shoot temperatures and transpiration. ? Methods: Root contraction was identified by measuring shoot descent and anatomical analysis. Temperatures and transpiration were measured for plants at two heights in sandy and rocky soil, and temperature tolerances were determined by vital staining. ? Key results: Plants embedded in rocky soil survived an extreme heat episode, unlike plants in sandy soil, though rocks did not moderate low temperatures. Root contraction occurred regardless of season and soil moisture. Xylem conduits (wide-band tracheids) formed a compressible lattice that decreased root length as rays enlarged the root base radially. Plant position in the soil did not affect transpiration. ? Conclusions: Contractile roots pulled plants of A. fissuratus into the soil at rates of 6-30 mm yr(-1). Maintaining shoots level with the soil surface kept plant temperatures below the high lethal temperature and improved survivorship in soil shaded by surface rocks.     

Root growth of subalpine and montane Eucalyptus seedlings at low soil temperatures

 This study examines the effect of different soil temperatures on root growth in seedlings of Eucalyptus pauciflora Sieber ex Sprengel subsp. pauciflora and Eucalyptus nitens (Deane & Maiden) Maiden. Seedlings were grown in a glasshouse in pots containing soil. Pots were held in water baths maintained at 3, 7 or 13°C, whilst shoots were exposed to ambient glasshouse temperatures. The experiments were designed to separate direct effects of soil temperature from effects due to differences in seedling size. In the first experiment, seedlings were grown to constant height (25 cm for both species), in the second to constant time (100 days for E. pauciflora and 64 days for E. nitens) and in the third experiment seedlings were transferred between soil temperatures. The rate of growth of both species increased with increasing soil temperature. E. nitens grew faster than E. pauciflora at 7 and 13°C, but E. pauciflora grew faster than E. nitens at 3°C. The rate of browning of roots increased with decreasing soil temperature and at a faster rate in E. nitens than E. pauciflora. Root length was highly correlated to root mass within diameter and colour classes (r² > 0.7). However, brown roots were heavier than white roots. Consequently, changes in root mass did not reflect changes in root length when the proportion of brown to white root also changed. For example, at a constant height of 25 cm at 3°C, E. nitens had greater root mass but lesser root length than E. pauciflora. E. pauciflora at 3°C grew faster, and had more root length and less brown roots than E. nitens. This supports the argument that E. pauciflora is better adapted than E. nitens to survive and grow at lower soil temperatures. Received: 16 December 1996 / Accepted: 2 April 1997     

Using a fine root extraction device to quantify small diameter corn roots (≥0.025 mm) in field soils

The study of fine roots growing under field conditions is limited by the techniques currently available for separating these roots from soil. This study had two objectives: to measure the total root length of field grown corn (Zea mays L.) by root diameter class, and to develop an inexpensive and efficient root washing device that would effectively capture all of the roots in a field soil sample. An inexpensive Fine Root Extraction Device (FRED) was constructed from readily available materials and was successful at extracting all roots, including very fine diameter roots (0.025 mm), from field soil samples. Greater than 99.7% of marked roots introduced to the FRED were recaptured by the device. Soil samples from three depths, and on three dates, from field grown corn were placed in the FRED. We found that more than 56% of total root length occurred in roots whose diameters were smaller than 0.175 mm, and more than 35% of root length occurred in roots smaller than 0.125 mm in diameter. Corn roots of the diameters described here have not been reported in field soils prior to this study. Root researchers who fail to measure these very fine roots will significantly underestimate root length density. Widespread use of the FRED should improve our understanding of root distribution in field soils.     

Growth of seedling roots in response to external osmotic stress by polyethylene glycol 20,000

Seedling roots of ten plant species were grown in siliceous sand wetted with solutions of polyethylene glycol (PEG) of MW=20,000 with osmotic potentials of 0.0, ? 0.25, ? 0.5 and ? 1.0 MPa. After 48 h growth under controlled lighting, root elongation and root diameter were measured. Root elongation of all species was reduced by increasing levels of external osmotic stress. Dicotyledonous species were affected more than monocotyledons at potentials of ? 0.25 and ? 0.5 MPa but less at ? 1.0 MPa. Root diameters of all the species were thicker than those of the unstressed at potentials of ? 0.25 and ? 0.5 MPa. At a potential of ? 1.0 MPa the dicotyledons were still thicker, though not by as much as they were at ? 0.25 and ? 0.5 MPa. The monocotyledons, in contrast, were thinner at ? 1.0 MPa. There was a significant positive correlation (r=0.81, p <0.01) between root diameter and root elongation at ? 1.0 MPa potential. Species were ranked according to the relative root elongation (RRE) and relative root thickness (RRT) at the highest level of stress (? 1.0 MPa). In both rankings dicotyledonous species were in the top ranks and monocotyledous species were in lower positions. The results are compared with those for the elongation and thickening of roots growing against external mechanical stress obtained in a previous study. There were good correlations between the responses observed for the two types of external stress. The implications of these findings are discussed.     

Comparison of the effects of nitrate and ammonium forms of nitrogen on auxin content in roots and the growth of plants under different temperature conditions

Average root length, root/shoot ratio and auxin content in roots were higher in plants supplied with nitrate rather than ammonium and grown at 18, 21, 24°C. The effects on root length were most pronounced at the highest temperatures (21 and 24°C); and the warmer the temperature, the earlier appearance of the differences in growth rate between NO₃- and NH₄-fed plants. A sharp acceleration of root growth was characteristic of NO₃-fed plants grown at 21 and 24°C and was associated with a temporary increase in auxin concentration measured by immunoassay.     

Biophysics of the growth responses of pea roots to changes in penetration resistance

Two experiments were performed in simplified soil-less systems to study how roots respond to changes in mechanical impedance. In the first the increases in root force and diameter that occur when a pea root was impeded mechanically inside a hole with rigid conical walls were determined. The experiment was performed at 8°C and at 25°C, and the root growth pressures generated were calculated during periods of 12, 24, 48 and 72 hours. The maximum growth pressures generated were approximately the same at both temperatures, although the maximum pressure was achieved approximately twice as quickly at 25°C than at 8°C, being reached within 15–20 hours. In the second set of experiments a new technique was developed to measure simultaneously the elongation rate and the force exerted by the roots of seedlings grown in moist air. A constant force was exerted by a force transducer on a pea radicle using a system of pulleys, and the elongation rate of the pea root was monitored using a linear variable differential transformer (LVDT). The changes in root elongation rate were recorded that occurred in response to increases and decreases in the applied force. Root elongation rate decreased by more than 50% within 30 min of increasing the applied force by 100 mN. A similarly fast, but smaller increase in growth rate occurred when the force was removed. The interpretation of results from both studies will be discussed in terms of a modified form of the Lockhart model of growth.     

A study of the role of root morphological traits in growth of barley in zinc-deficient soil

Zinc (Zn) deficiency reduces crop yields globally. This study investigated the importance of root morphological traits, especially root hairs, in plant growth and Zn uptake. Wild-type barley (Hordeum vulgare) Pallas and its root-hairless mutant brb were grown in soil and solution culture at different levels of Zn supply for 16 d. Root morphological traits (root length, diameter, and surface area) were measured using the WinRHIZOPro Image Analysis system. In soil culture, Pallas had greater shoot dry matter, shoot Zn concentration, shoot Zn content, and Zn uptake per cm(2) root surface area than brb, primarily under zinc deficiency. Both Pallas and brb developed longer roots under Zn deficiency. Development of root hairs was not affected by plant Zn status. In solution culture, there were no significant genotypic differences in any of the parameters measured, indicating that mutation in brb does not affect growth and Zn uptake. However, both Pallas and brb developed longer and thinner roots, and root hair growth was less than in soil culture, and was not affected by plant Zn status. The better growth and greater Zn uptake of Pallas compared with brb in Zn-deficient soil can be attributed primarily to greater root surface area due to root hairs in Pallas rather than other root morphological differences.     

Effects of temperature on the development and growth of winter wheat roots

Wheat plants were grown in columns of soil until early stem elongation at a wide range of constant root temperatures. Two light environments were imposed and three levels of nitrogen fertilizer added at sowing. Shoot and root development and growth were measured by destructive sampling to investigate the combined effects of temperature and changing nutrient and assimilate supply. Both mainstem leaf and root axis production were linearly related to thermal time above a base temperature of 0°C. Low irradiance affected the appearance of mainstem tillers and associated nodal root axes. Nitrogen had little effect on shoot or root development but increased shoot area between 6 and 8 mainstem leaves. Higher temperatures and supplementary light resulted in larger root systems when compared at equivalent times after sowing. Total root length and root dry weight increased exponentially with thermal time, based on the mean of 4 cm soil and 2 cm air temperatures, but no single relation existed for all temperature and light treatments. Total plant dry matter, root length and root dry weight increased linearly with accumulated, intercepted, photosynthetically active radiation. Root growth responded less than the shoot to supplementary light. Increasing temperature reduced the proportion of root weight to total plant weight.     

Seasonal variation of growth and development of the roots of five second year conifer species in the nursery

Every other week over their second growing season, stem height, collar diameter, shoot and root dry masses, number of lateral roots and length of the tap root were measured on nursery grown seedlings ofAbies balsamea L. Mill.,Pinus banksiana Lamb.,Pinus resinosa Ait.,Picea mariana Mill. BSP andPicea glauca Moench Voss. Root elongation, branching and mycorrhizal development were also recorded.Given species showed distinct seasonal growth patterns. The rate and timing of maximum root growth (mg/dry weight/week) differed markedly between species.Except for the increase in height ofPinus banksiana, root and shoot growth were not negatively correlated.The results are discussed in relation to the performance of tree seedlings in the nursery.     

Ethylene Evolution from Maize (Zea mays L.) Seedling Roots and Shoots in Response to Mechanical Impedance

The effect of mechanical impedance on ethylene evolution and growth of preemergent maize (Zea mays L.) seedlings was investigated by pressurizing the growth medium in triaxial cells in a controlled environment. Pressure increased the bulk density of the medium and thus the resistance to growth. The elongation of maize primary roots and preemergent shoots was severely hindered by applied pressures as low as 10 kilopascals. Following a steep decline in elongation at low pressures, both shoots and roots responded to additional pressure in a linear manner, but shoots were more severely affected than roots at higher pressures. Radial expansion was promoted in both organs by mechanical impedance. Primary roots typically became thinner during the experimental period when grown unimpeded. In contrast, pressures as low as 25 kilopascals caused a 25% increase in root tip diameter. Shoots showed a slight enhancement of radial expansion; however, in contrast to roots, the shoots increased in diameter even when growing unimpeded. Such morphological changes were not evident until at least 3 hours after initiation of treatment. All levels of applied pressure promoted ethylene evolution as early as 1 hour after application of pressure. After 1 hour, ethylene evolution rates had increased 10, 32, 70, and 255% at 25, 50, 75, and 100 kilopascals respectively, and continued to increase linearly for at least 10 hours. When intact corn seedlings were subjected to a series of hourly cycles of pressure, followed by relaxation, ethylene production rates increased or decreased rapidly, illustrating tight coupling between mechanical impedance and tissue response. Seedlings exposed to 1 microliter of ethylene per liter showed symptoms similar to those shown by plants grown under mechanical impedance. Root diameter increased 5 times as much as the shoot diameter. Pretreatment with 10 micromolar aminoethoxyvinyl glycine plus 1 micromolar silver thiosulfate maintained ethylene production rates of impeded seedlings at basal levels and restored shoot and root extension to 84 and 90% of unimpeded values, respectively. Our results support the hypothesis that ethylene plays a pivotal role in the regulation of plant tissue response to mechanical impedance.