Effect of soil moisture and phosphate level on root hair growth of corn roots

Summary Root hairs have been shown to enhance P uptake by plants growing in low P soil. Little is known of the factors controlling root hair growth. The objective of this study was to investigate the influence of soil moisture and P level on root hair growth of corn (Zea mays L.). The effect of volumetric soil moistures of 22% (M₀), 27% (M₁), and 32% (M₂) and soil (Raub silt loam, Aquic Argiudoll) P levels of, 0.81 (P₀), 12.1 (P₁), 21.6 (P₂), 48.7 (P₃), and 203.3 (P₄) mol P L^–1 initially in the soil solution, on shoot and root growth, P uptake, and root hair growth of corn was studied in a series of pot experiments in a controlled climate chamber. Root hair growth was affected more by soil moisture than soil P. The percentage of total root length with root hairs and the density and length of root hairs on the root sections having root hairs all increased as soil moisture was reduced from M₂ to M₀. No relationship was found between root hair length and soil P. Density of root hairs, however, was found to decrease with an increase in soil P. No correlation was found between root hair growth parameters and plant P content, further suggesting P plays a secondary role to moisture in regulating root hair growth in soils. The increase in root hair growth appears to be a response by the plant to stress as yield and P uptake by corn grown at M₀ were only 0.47 to 0.82, and 0.34 to 0.74, respectively, of that measured at M₁ across the five soil P levels. The increase in root hair growth at M₀, which represents an increase of 2.76 to 4.03 in root surface area, could offset, in part, the reduced rate of root growth, which was the primary reason for reduced P uptake under limited soil moisture conditions.Journal Paper No. 10,066 Purdue Univ. Agric. Exp. Stn., W. Lafayette, IN 47907. Contribution from the Dep. of Agron. This paper was supported in part by a grant from the Tennessee Valley Authority.     

Transformation and mineralization of soil organic nitrogen by the humivorous larva of Pachnoda ephippiata (Coleoptera: Scarabaeidae)

In common bean (Phaseolus vulgaris L.), Fusarium root rot (caused by Fusarium solani f. sp. phaseoli) disease severity is increased by environmental factors that stress the plant. The current study used reciprocal grafting techniques with the resistant cultivar FR266 and the susceptible cultivar Montcalm to determine if the genetic control of resistance is conferred by the rootstock (root genotype) or the scion (shoot genotype) and if root vigor played a role in resistance. The influence of a compacted layer on root and shoot genotype response and root rot resistance was studied. Root rot resistance was found to be controlled by the root genotype, such that on a scale of 1 to 7 (severe disease) the FR266 root had an average score of 2.3 and the Montcalm root had an average score of 4.4. However, when grafted plants were grown in the presence of a compacted layer, the FR266 root and/or shoot genotype in any graft combination with the susceptible Montcalm had reduced root rot (score = 2.4 average) than the Montcalm self graft (score = 4.5). Root mass was shown to be controlled by the root genotype in the absence of compaction such that the FR266 root was 26% larger that the Montcalm root when grafted onto a FR266 shoot or a Montcalm shoot. When a compacted layer was present the root and shoot genotype both contributed to root mass. Average root diameter was controlled by the shoot genotype, as the FR266 shoot grafted to Montcalm or FR266 roots had thicker roots (average diameter 0.455 mm) than the Montcalm shoot (average diameter 0.418 mm). This study shows evidence that root vigor in the presence of Fusarium disease pressure should be evaluated to effectively develop common bean lines resistant to Fusarium root rot across a range of environments.     

Root water uptake and profile soil water as affected by vertical root distribution

Water uptake by plant roots is a main process controlling water balance in field profiles and vital for agro-ecosystem management. Based on the sap flow measurements for maize plants (Zea mays L.) in a field under natural wet- and dry-soil conditions, we studied the effect of vertical root distribution on root water uptake and the resulted changes of profile soil water. The observations indicate that depth of the most densely rooted soil layer was more important than the maximum rooting depth for increasing the ability of plants to cope with the shortage of water. Occurrence of the most densely rooted layer at or below 30-cm soil depth was very conducive to maintaining plant water supply under the dry-soil conditions. In the soil layers colonized most densely by roots, daytime effective soil water saturation (S _e) always dropped dramatically due to the high-efficient local water depletion. Restriction of the rooting depth markedly increased the difference of S _e between the individual soil layers particularly under the dry-soil conditions due likely to the physical non-equilibrium of water flow between the layers. This study highlights the importance of root distribution and pattern in regulating soil water use and thereby improving endurance of plants to seasonal droughts for sustainable agricultural productivity.     

Root characteristics of representative Mediterranean plant species and their erosion-reducing potential during concentrated runoff

Gully erosion is an important soil degradation process in Mediterranean environments. Revegetation strategies for erosion control rely in most cases on the effects of the above-ground biomass on reducing water erosion rates, whereas the role of the below-ground biomass is often neglected. In a Mediterranean context, the above-ground biomass can temporally disappear because of fire or overgrazing and when concentrated flow erosion occurs, roots can play an important role in controlling soil erosion rates. Unfortunately, information on root characteristics of Mediterranean plants, growing on semi-natural lands, and their effects on the topsoil resistance to concentrated flow erosion is lacking. Therefore, typical Mediterranean grass, herb, reed, shrub and tree root systems of plants growing in habitats that are prone to concentrated flow erosion (i.e. in ephemeral channels, abandoned fields and steep badland slopes) are examined and their erosion-reducing potential was evaluated. Root density (RD), root length density (RLD) and root diameters are measured for 26 typical Mediterranean plant species. RD values and root diameter distribution within the upper 0.10–0.90 m of the soil profile are then transformed into relative soil detachment rates using an empirical relationship in order to predict the erosion-reducing effect of root systems during concentrated runoff. Comparing the erosion-reducing potential of different plant species allows ranking them according to their effectiveness in preventing or reducing soil erosion rates by concentrated flow. RD in the 0.10 m thick topsoil ranges between 0.13 kg m⁻³ for Bromus rubens (L.) and 19.77 kg m⁻³ for Lygeum spartum (L.), whereas RLD ranges between 0.01 km m⁻³ for Nerium oleander (L.) and 120.43 km m⁻³ for Avenula bromoides ((Gouan) H. Scholz.) Relative soil detachment rates, compared to bare soils, range between 0.3 × 10^-12 and 0.7 for the 0.10 m thick topsoil. The results show that grasses such as Helictotrichon filifolium ((Lag.) Henrard), Piptatherum miliaceum ((L.) Coss.), Juncus acutus (L.), Avenula bromoides ((Gouan) H. Scholz), Lygeum spartum (L.) and Brachypodium retusum ((Pers.) Beauv.) have the highest potential to reduce soil erosion rates by concentrated flow in the 0–0.1 m topsoil. But also shrubs such as Anthyllis cytisoides (L.) and Tamarix canariensis (Willd.), having high root densities in the topsoil, can reduce erosion rates drastically. Among the species growing in channels, Juncus acutus (L.) has the highest erosion reducing potential, whereas Phragmites australis (Cav.) is the least effective. On abandoned fields, Avenula bromoides ((Gouan) H. Scholz) and Plantago albicans (L.) are the most effective species in reducing concentrated flow erosion rates, while Thymelaea hirsuta (L. (Endl.)) and Bromus rubens (L.) perform the worst. On steep badland slopes, Helictotrichon filifolium ((Lag.) Henrard) and Anthyllis cytisoides (L.) perform the best in the analysis of erosion reducing potential, while Ononis tridentata (L.) is the least effective species. These findings have implications for ecological restoration and management of erosion-prone slopes.     

Root growth and water uptake in winter wheat under deficit irrigation

Root growth is critical for crops to use soil water under water-limited conditions. A field study was conducted to investigate the effect of available soil water on root and shoot growth, and root water uptake in winter wheat (Triticum aestivum L.) under deficit irrigation in a semi-arid environment. Treatments consisted of rainfed, deficit irrigation at different developmental stages, and adequate irrigation. The rainfed plots had the lowest shoot dry weight because available soil water decreased rapidly from booting to late grain filling. For the deficit-irrigation treatments, crops that received irrigation at jointing and booting had higher shoot dry weight than those that received irrigation at anthesis and middle grain filling. Rapid root growth occurred in both rainfed and irrigated crops from floral initiation to anthesis, and maximum rooting depth occurred by booting. Root length density and dry weight decreased after anthesis. From floral initiation to booting, root length density and growth rate were higher in rainfed than in irrigated crops. However, root length density and growth rate were lower in rainfed than in irrigated crops from booting to anthesis. As a result, the difference in root length density between rainfed and irrigated treatments was small during grain filling. The root growth and water use below 1.4 m were limited by a caliche (45% CaCO₃) layer at about 1.4 m profile. The mean water uptake rate decreased as available soil water decreased. During grain filling, root water uptake was higher from the irrigated crops than from the rainfed. Irrigation from jointing to anthesis increased seasonal evapotranspiration, grain yield, harvest index and water-use efficiency based on yield (WUE), but did not affect water-use efficiency based on aboveground biomass. There was no significant difference in WUE among irrigation treatments except one-irrigation at middle grain filling. Due to a relatively deep root system in rainfed crops, the higher grain yield and WUE in irrigated crops compared to rainfed crops was not a result of rooting depth or root length density, but increased harvest index, and higher water uptake rate during grain filling.     

Root distribution and water uptake patterns of maize cultivars field-grown under differential irrigation

Summary Rooting and water uptake patterns were determined for three maize (Zea mays L) varieties field-grown during the 1983/84 dry season under seven irrigation levels on a sandy loam soil. Roots were mainly concentrated in the top 22 cm due to a 40 cm thick compact gravelly layer occurring from about this depth in the profile. There were significant varietal differences, distinguished by root length density (RLD) and length/weight ratio (LAR) distributions at depth and at varying soil moisture regimes. These properties were related to water extraction patterns and grain yields. Yields obtained at adequate soil moisture were 6.9 tha⁻¹ for TZESR-W (var 1), 4.2 t/ha for TZSR-W (var 2) and 3.7t ha⁻¹ for FARZ-7 (var 3). These yeilds were respectively associated with maximum RLD of 2.56, 1.88 and 1.70 cm cm⁻³ and corresponding LWR of 2.64, 1.93 and 1.62 cm mg⁻¹. Average seasonal water uptake was estimated at 4.2, 3.0 and 2.8 mm day⁻¹ for var 1, 2 and 3, respectively. Better performance of var 1 was attributed to the development of a more active and deep rooting system.     

大田期烟草根系构型参数的动态变化

采用“根箱”法研究了大田期烟草根系构型参数在时间、空间上的动态变化.结果表明, 烟草2级侧根总长度的增加明显大于1级侧根,根快速增长期分别出现在移栽后26～40和56～70 d.栽后57 d(打顶)前,烟草根系的分枝密度表现为10～20＞0～10＞20～30＞30～40 cm,此后随土层的加深呈递减趋势.在主根上,以7～21 cm范围内的分枝密度最大.打顶前,比根长随着入土深度的加深而递增;栽后90 d,比根长随土层的加深而递减.1级侧根根长密度在0～10 cm土层内的变化呈“S”型曲线,10～20、20～30和30～40 cm内表现为双峰曲线;2级侧根根长密度随生育期的进程而增加,其中0～10 cm根长密度的变化为“S”曲线,其它层次为单峰曲线.     

黄土高原土壤紧实度对蚕豆生长的影响

通过盆栽试验、连续 2年的田间小区试验和农户生产试验 ,研究了土壤紧实状况对蚕豆 (Viciafa ba)生长的影响 ,讨论了当地土壤容重较高的原因 ,并提出了改进措施 .结果表明 ,随着 0～ 7cm土层土壤容重的增加 ,蚕豆植株每株的茎与根干重降低 ,根腐病 (Fusariumspp .)引起的死亡率增加 ,种子产量减少 .田间试验条件下 ,与生长于容重为 1.5 5和 1.6 4 g·cm-3 小区内的植株相比 ,生长于容重 1.84 g·cm-3 小区内的植株每株茎与根干重可分别减少 2 7.9%和 30 .8% ,植株累计死亡率增加 2 1.0 %～ 4 8.7% ,种子产量每公顷减少 19.8% .在 8户蚕豆田中进行的多点生产试验表明 ,春季土壤容重与蚕豆幼苗的根与茎干重、秋季土壤容重与种子产量均呈显著负相关     

水稻根系生长及其对土壤紧密度的反应

采用筒栽方法研究了杂交稻汕优63和65002在分蘖期,穗分化期,开花期和成熟期根系生长量和垂直分布以及开花期土壤容重对根系生长和分布的影响,结果表明,根系生长量以开花期最高,开花后下降,随生育进展,深层根系（20-45cm)比例提高,分蘖期到穗 分化期是根系重量增长最快,且根系向下生长的主要时期,随耕层以下土壤容重提高,根系生长量下降,同时深层根系比例下降。     

Estimating root lifespan of two grasses at contrasting elevation in a salt marsh by applying vitality staining on roots from in-growth cores

Contrasting soil conditions caused by different inundation frequenciesrequire different root growth strategies along the elevational gradient ofcoastal salt marshes. The objective of this study was to examine (1) if rootlifespan was shorter in Elymus pycnanthus, a relativelyfast-growing competitive species dominating high marshes, than inSpartina anglica, a relatively slow-growingstress-tolerating species dominating low marshes, and (2) if the species withlonger lifespan had higher tissue density (g cm^–3) and lowerspecific root length (m g^–1) than the species with shorterlifespan. Root production and mortality rates were established by samplingrootsin in-growth cores, and using triphenyltetrazolium chloride (TTC) staining todistinguish vital from dead roots. Root lifespan was estimated by dividing theliving root biomass (Elymus: 36 gm^–2, Spartina: 100 gm^–2) by root production (Elymus:0.28 g day^–1 m^–2,Spartina: 0.25 g day^–1m^–2) or root mortality rates(Elymus: 0.42–0.53 g day^–1m^–2). Spartina did not exhibitsubstantial mortality. Despite the present method only yielding rough estimatesof average root lifespan, it is evident that root longevity is much shorter inElymus than in Spartina. Rootlifespanranged between 10–19 weeks for Elymus but was closeto 1 year in Spartina, indicating thatElymus replaces it's roots continuously throughout thegrowing season, whereas Spartina maintains its roots overthe growing season. Fine roots of Elymus had slightlylowertissue density (0.094) than those of Spartina (0.139),whereas coarse roots of Elymus andSpartina had similar tissue density (0.100 gcm^–3). Fine roots of Elymus andSpartina had similar specific root length (195 mg^–1). However, coarse roots ofElymus (50 m g^–1) had higherspecific root length than those of Spartina (20 mg^–1) due to having smaller root diameter(Elymus: 548 m,Spartina: 961 m). We conclude thatpresentobservations on Elymus and Spartinasupport our first hypothesis that the competitive species fromthehigh marsh had short-lived roots compared to the'stress-tolerating'species from the low marsh. However, our result provide only weak support forthe existence of a positive correlation between root longevity and tissuedensity and a negative correlation between root longevity and specific rootlength.     

Effect of soil volume and plant density on mycorrhizal infection and growth response

Summary The effect of soil volume and plant density on mycorrhizal infection and growth response was studied with onion. There was a significant negative correlation between percentage vesicular-arbuscular mycorrhizal infection and root density. The growth response due to mycorrhiza decreased when less soil was available for the plant. The root: shoot ratio decreased with increasing plant density in both mycorrhizal and non-mycorrhizal plants. Pot size did not affect the root: shoot ratio.     

Exploring the interacting effect of soil texture and bulk density on root system development in tomato (Solanum lycopersicum L.)

Knowledge of the responses of root systems in horizoned heterogeneous soil is vital to optimise uptake of water and nutrients to maximise crop productivity. We explored the interacting effects of soil bulk density and texture on the development of root systems in tomato.Two main techniques were employed, X-ray micro-Computed Tomography (μCT), to provide non-destructive, three-dimensional (3D) images of root systems in situ and destructive root washing followed by WinRHIZO^® scanning. Solanum lycopersicum L. cv. Ailsa Craig plants were grown in soil columns for 10 days to measure the effect of soil compaction on selected root traits. Treatments included bulk density (1.2–1.6 Mg m⁻³), soil texture (loamy sand and clay loam) and the effects of layering.The effect of bulk density on root growth was greatest 3 days after transplanting (DAT) in both soil types. The effect of soil texture was not apparent at this stage, but was significant at 10 DAT for most root and shoot variables. The influence of bulk density differed between soil types as increasing compaction promoted plant growth in clay loam but retarded root growth in loamy sand.We observed that at 3 DAT root growth is primarily influenced by bulk density but by 10 DAT a switch in the processes regulating root growth occurs and the texture of the soil becomes very influential. Future investigations of root growth must consider soil physical properties individually and at specific time points, as their importance changes as the root system becomes established. Here we have demonstrated both positive and negative impacts across a wide range of bulk density treatments in different soil textures on root growth. This illustrates the importance of understanding the complex nature of root–soil interactions, especially for agricultural practices such as seedbed preparation.     

Vertical distribution of fine roots in relation to soil factors in Pinus tabulaeformis Carr. forest of the Loess Plateau of China

Growth and vertical distribution of fine root closely depend on soil resource availability. Better understanding of relationships of root profile with vertical distribution of available soil resource and soil characteristics can allow ecologists to predict the fine root distribution on the scales ranging from individual plants to vegetation communities. The objective of the study was to understand the fine root mass density (FRMD), fine root length density (FRLD), fine root area density (FRAD), mean root diameter and specific root length (SRL), vertical distribution in soil profile and their relation with soil environment factors in semiarid and arid Loess Plateau of China. The vertical fine root distribution and soil bulk density, soil moisture and soil inorganic N in 0-60 cm soil profile (0–15, 15–30, 30–45 and 45–60 cm intervals) were investigated by soil coring methods in three Pinus tabulaeformis Carr. forests chosen at three locations. The fine root density parameters (FRMD, FRLD and FRAD) and SRL peaked in the most upper soil layer (0–15 cm interval) and decreased with increased soil depth. The results provided a strong support that soil water rather than soil inorganic N is a key control on fine root distribution in the Loess Plateau. With increased soil moisture, the root mass, length and SRL increased and the mean root diameter decreased. The effects of soil bulk density on the fine root parameters were consistent with those of the soil water. An unexpected result was obtained about the relationships between soil organic N and the root distributions and occurrences because of no differences among the soil depth intervals in soil inorganic N content. It might be associated with severe soil water deficit limiting soil nitrogen utilization efficiency in arid Loess Plateau.     

转基因作物对土壤生态系统的影响

综述了转基因作物对土壤生态系统影响的研究进展,包括转基因作物中的外源基因在土壤中的活性,转基因作物对土壤微生物区系有土壤酶活性的影响以及转基因作物对土壤动物区系的影响,转基因作物对土壤生态系统的影响与导入的外源基因特性和土壤类型相关,转基因产物进入土壤后引起的土壤生物变化的程度依赖于许多因素,最重要的决定因素是生态系统的复杂性和稳定性,评价不同转基因作物对土壤生态系统的影响具有重要的生态学意义,急需发展和完善以分子生物学为主的风险评价方法。     

The response of sorghum and sunflower to short-term waterlogging

Summary Sorghum and sunflower were waterlogged for nine days during the vegetative, floral initiation/buds-visible or anthesis stage of growth under glasshouse conditions to observe the effects on root growth and development. In addition, some plants were waterlogged at all three stages to observe any adaptations induced by waterlogging. The most marked effects occurred at the initiation/buds-visible stage where a 30% reduction in root length and a 40% reduction in root dry weight of sorghum occurred with comparable figures for sunflower being 50 and 60% respectively. Generally, sorghum roots had a higher porosity than sunflower which may contribute to its greater tolerance to waterlogging. The observed changes in root growth are discussed in relation to previously documented effects of waterlogging on growth of the two species and changes which occur in the soil environment.     

Root growth and distribution of two short-season rice genotypes

Root growth dynamics of lowland rice (Oryza sativa L.) throughout the growing season are poorly understood. A field experiment was conducted in 1987 to compare root growth and distribution of two rice genotypes at two Arkansas locations on soils with different physical and chemical properties. Two genotypes, Bond and an experimental line (RU8701084), were grown on a Captina silt loam (Typic Fragiudults) at Fayetteville, AR, and on a Crowley silt loam (Typic Albaqualfs) near Stuttgart, Ar. Plots contained minirhizotrons oriented at a 45° angle and extended 55 cm (Captina) and 40 cm (Crowley) vertical to the soil surface. Root measurements were taken several times during the season at specific growth stages. Plant height and tiller number were taken at 9 dates at Fayetteville up to physiological maturity. In general, root length (RL) and root length density (RLD) were greater on the Captina soil. Genotypes at both locations reached maximum root growth rates between active tillering and panicle initiation (PI) and maximum RL by early reproduction. Total RL were similar between genotypes on the Captina. However on the Crowley, the mean RL for Bond between the period of early booting and flood removal was an average of 54% greater than for RU8701084. During early reproductive growth at both locations RL plateaued, but then declined during the grain filling process. There was a trend for RU8701084 to contain a greater percentage of its total RL in the top 20 and 10 cm of soil on the Captina and Crowley, respectively, while Bond tended to be a deeper rooted genotype. Bond had a greater RLD at the 20–30 cm depth increment on the Crowley, which contributed to the greater RL. Less than 15% of the total RL of either genotype was measured below 30 cm on the Crowley. In contrast, nearly 25% of the total RL was found at the 30–40 cm depth increment on the Captina. Results showed that rice root growth varied between soils, that root distribution patterns differed between genotypes, and that patterns of root growth changed with changes in plant development.     

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     

Effect of soil acidity and saturating cation on adsorption of urea in soil

Summary Elevating the pH of two extremely acid tropical soils from an initial pH of 4 to 5 with Ca(OH)₂ and NaOH solutions resulted in a sharp decrease in urea adsorption. Further increases in pH to 9 caused only slight further decrease in urea adsorption. Ca treatment resulted in slightly higher adsorption than Na treatment over the full range of pH values studied for a clay soil with high smectite content, and for a loam with kaolinite mineralogy below pH 7. Above pH 7 a kaolinitic loamy soil gave slightly higher adsorption with Na treatment than with Ca.     

Effect of soil compaction on growth, yield and light interception of selected crops

The response of spring barley (Hordeum vulgare, cvs Carnival and Atem), faba beans (Vicia faba, cv. Maris Bead), sugar beet (Beta vulgaris, cv. Monoire), forage maize (Zea mays, cv. Leader), forage peas (Pisum sativum, cv. Poneka) and white turnip (Brassica campestris, cv. Barkant) to topsoil compaction was investigated in a three year trial. Soil compaction was induced by tractor wheeling after crop sowing. Compaction reduced leaf area and dry matter accumulation in all crops in every season. Yield of barley was reduced by 29%, 27% and 40% in 1984, 1986 and 1987 respectively. Yield of maize, peas and turnip decreased by 33%, 14% and 13% in 1986 and 25%, 16% and 19% in 1987. Yields of beans and sugar beet were decreased by 34% and 35% respectively in 1984. Light interception was decreased in all crops in all three years of study but, with the exception of maize in 1987, the efficiency of conversion of radiant energy to dry matter was not significantly affected by soil compaction. It is concluded that reduced dry matter production and yield due to soil compaction was more a consequence of reduced light interception because of restricted leaf area development rather than as a result of an impaired ability of crops to utilise intercepted radiant energy.