Rate of differentiation and emergence of nodal maize roots

The timing of root production is one of the parameters required for modelling the root system architecture. The objectives of this study are (1) to describe the rate of appearance of adventitious root primordia of maize and their rate of emergence out of the stem; (2) to test equations for the prediction of the rank of the phytomer on which root emergence occurs, in a wide range of field situations.Maize, cultivar Dea, was grown in controlled conditions and in the field in 1987, 1988, 1989 and 1991. Plants were regularly sampled and the following data were recorded: foliar stage, number of root primordia and number of emerged roots per phytomer. Root primordia were counted in transverse thin sections in the stem.At a single plant level, root primordia differentiation occurred sequentially on the successive phytomers, with no overlapping between two phytomers. The same was true for root emergence. Roots belonging to the same phytomer emerged at approximately the same time.At a plant population level, there was a linear relationship between the rank of the phytomer on which root primordia were differentiated and cumulated degree-days after sowing. A linear relationship was also observed between the rank of the phytomer on which roots were emerging and cumulated degree-days or foliar stage. In the range of field situations tested (several years, sowing dates and planting densities), both equations gave an accurate prediction of the timing of root emergence during the plant cycle.     

A developmentally based categorization of branching in Trifolium repens L.: influence of nodal roots

This study describes the successive stages of development of branches from axillary buds in fully rooted plants of Trifolium repens grown in near optimal conditions, and the way in which this developmental pathway differs when nodal root formation is prevented as plants grow out from a rooted base. Cuttings of a single genotype were established in a glasshouse with nodal root systems on the two basal phytomers and grown on so that nodal rooting was either permitted (+R) or prevented (-R). In +R plants, axillary tissues could be assigned to one of four developmental categories: unemerged buds, emerged buds, unbranched lateral branches or secondarily branched lateral branches. In -R plants, branch development was retarded, with the retardation becoming increasingly pronounced as the number of -R phytomers on the primary stolon increased. Retarded elongation of the internodes of lateral shoots on -R plants resulted in the formation of a distinct fifth developmental category: short shoots (defined as branches with two or more leaves but with mean internode length equal to, or less than, 10% of that of the immediately proximal internode on the parent stolon) which had reduced phytomer appearance rates but retained the potential to develop into lateral branches. Transfer of +R plants to -R conditions, and vice versa, after 66 d demonstrated that subsequent branch development was wholly under the control of the youngest nodal root present, regardless of the age and number of root systems proximal to it.     

The relationships between static and dynamic variables in the description of root growth. Consequences for field interpretation of rooting variability

Field root investigations are often limited by the static nature of classical observations, resulting in the need to develop alternative methodologies that allow dynamic interpretation of root architecture variability on the basis of static measurements. The objectives of this work were (i) to evaluate the use of selected morphological indicators, namely root apical diameter (Da) and the length of the apical unbranched zone (LAUZ), in predicting primary and lateral root growth patterns in banana trees, (ii) to propose a field methodology for the assessment of root dynamics based on static measurements. Banana trees (Musa acuminata cv `Grande Naine') were grown in 5 rhizotrons as well as in field conditions, respectively on pouzzolane and Mollic Andosols. In rhizotrons, root growth analysis was carried out by reporting root elongation, Da and LAUZ, three times a week. In field conditions, 4 series of excavations were made at three-week intervals. Apart from root growth rate, measurements were the same as those in the rhizotrons. LAUZ was confirmed as a stable and good predictor of root growth rate for the different types of roots. In the rhizotrons, the root growth of lateral roots was found to be well correlated to the product of Da and the growth rate of the bearing root. Evaluation in field conditions from static observations attested consistent relationships between measured and predicted root length for lateral roots (slopes close to 1:1). The apical diameter can be considered as a good indicator of root growth potential, while actual lateral root growth depends on the bearing root elongation rate. Morphological static indicators calibrated from growth dynamics in rhizotrons are of major interest in explaining growth variability in field conditions. Especially the `growth rate-LAUZ' relationship can be considered a useful tool in interpreting field patterns of growing roots in relation to various soil conditions.     

Effect of mutual shading on the emergence of nodal roots and the root/shoot ratio of maize

The effect of mutual shading on the root/shoot ratio and on the number of nodal roots of maize was studied. Plants of two varieties (Dea and LG2281) were grown in individual pots of 9 L, at three plant densities: 7.5, 11 and 15 plants m^–2. A control experiment was carried out in order to study if root growth was affected by the small size of the pots. Maize plants (cv Dea) were grown at a low plant density (7.5 plants m^–2) in pots of two different volumes (9 and 25 L respectively). In both experiments plants were watered every two hours with a nutrient solution. Some plants were sampled at five dates in the main experiment and the following data were recorded: foliar stage; root, stem and leaf dry weight; number of root primordia and number of emerged roots per phytomer. The final sampling date occurred at silking.Results of the control experiment showed that the root biomass was lower in small pots but the number of nodal roots per phytomer was not affected.Results of the main experiment showed that the total plant biomass and the root/shoot ratio were lower at high plant density. The number of emerged roots was strongly reduced on the upper phytomer (P₈). This reduction was mainly due to a lower percentage of root primordia which elongated. A proposed interpretation is that the number of roots which emerge on upper phytomers is controlled by carbohydrate availability.     

A comprehensive analysis of root morphological changes and nitrogen allocation in maize in response to low nitrogen stress

The plasticity of root architecture is crucial for plants to acclimate to unfavourable environments including low nitrogen (LN) stress. How maize roots coordinate the growth of axile roots and lateral roots (LRs), as well as longitudinal and radial cell behaviours in response to LN stress, remains unclear. Maize plants were cultivated hydroponically under control (4 mm nitrate) and LN (40 μm ) conditions. Temporal and spatial samples were taken to analyse changes in the morphology, anatomical structure and carbon/nitrogen (C/N) ratio in the axile root and LRs. LN stress increased axile root elongation, reduced the number of crown roots and decreased LR density and length. LN stress extended cell elongation zones and increased the mature cell length in the roots. LN stress reduced the cell diameter and total area of vessels and increased the amount of aerenchyma, but the number of cell layers in the crown root cortex was unchanged. The C/N ratio was higher in the axile roots than in the LRs. Maize roots acclimate to LN stress by optimizing the anatomical structure and N allocation. As a result, axile root elongation is favoured to efficiently find available N in the soil.     

Relationship between root elongation rate and diameter and duration of growth of lateral roots of maize

The objective of this work was to describe the relationship between elongation rate and diameter of maize roots and to estimate the length and growth duration of lateral roots of maize. Diameters and elongation rates of roots were measuredin situ on plants grown 5 weeks in small rhizotrons under greenhouse conditions. At the end of the experimental period the roots were harvested and diameters of axile and lateral roots were measured. The frequency distribution of diameters of harvested roots was bimodal with a minimum at 0.6 mm; 97% of axile roots were larger than this value and 98% of the lateral roots were smaller. Root elongation per day increased as diameter increased but the slope of the relationship with lateral roots was about 2.5 times that with axile roots when separate linear regressions were fitted to the two populations. The length of lateral roots found on axillary roots between the base and about 30 cm from the apex was approximately 2.2 cm. All of the data was consistent with the hypothesis that the lateral roots grew for about 2.5 days and then ceased growing. The axillary roots continued to grow throughout the experimental period at a rate of about 3 cm day⁻¹. Contribution from the Department of Agronomy, New York State College of Agriculture and Life Sciences, Cornell University, Ithaca, NY 14853. Agronomy paper No. 1661. This research is part of the program of the Center for Root-Soil Research.     

Evaluation in field conditions of a three-dimensional architectural model of the maize root system: Comparison of simulated and observed horizontal root maps

Most existing water and nutrient uptake models are based on the assumption that roots are evenly distributed in the soil volume. This assumption is not realistic for field conditions, and significantly alters water or nutrient uptake calculations. Therefore, development of models of root system growth that account for the spatial distribution of roots is necessary.The objective of this work was to test a three dimensional architectural model of the maize root system by comparing simulated horizontal root maps with observed root maps obtained from the field. The model was built using the current knowledge on maize root system morphogenesis and parameters obtained under field conditions. Simulated root maps (0.45 × 0.75 m) of horizontal cross sections at 3 depths and 3 dates were obtained by using the model for a plant population. Actual root maps were obtained in a deep, barrier-free clay-loamy soil by digging pits, preparing selected horizontal planes and recording root contacts on plastic sheets.Results showed that both the number of cross-sections of axile roots, and their spatial distribution characterized with the R-index value of Clark and Evans (1954), were correctly accounted for by the model at all dates and depths. The number of cross-sections of laterals was also correctly predicted. However, laterals were more clustered around axile roots on simulated root maps than on observed root maps. Although slight discrepancies appeared between simulated and observed root maps in this respect, it was concluded that the model correctly accounted for the general colonization pattern of the soil volume by roots under a maize crop.     

Dynamics of the Elongation of Internodes in Maize (Zea mays L.): Analysis of Phases of Elongation and their Relationships to Phytomer Development

The kinetics of elongation of individual internodes of maizestems were studied under field conditions. Thermal time coursesof internode length were recorded using non-destructive methods,based on direct measurement of X-ray photographs or on indirectestimates using heights of leaf collars. These data were complementedby serial dissections of maize stems, and by precise observationof the process of sheath emergence, to specify its role in thekinetics of internode elongation. The kinetics of elongationwere found to be composed of four phases. The rate of elongationrises exponentially during phase I, and then increases sharplyduring a short period (phase II), which is followed by a majorperiod of constant growth rate (phase III) and a shorter periodin which the rate declines (phase IV). During phase I, elongationappears to be integrated at the level of the whole apical cone.From phase II onwards, elongation becomes determined at thelevel of the phytomer. The emergence of the sheath attachedto the internode appears to be a possible trigger for the transitionbetween phase I and phase II, and it may also be involved invariation in final length among phytomers. Copyright 2000 Annalsof Botany Company Zea mays L., internode, elongation, modelling, dynamics, X-rays, collar, phytomer, stem, thermal time, phasic development     

Effect of light quality (red: far-red ratio) and defoliation treatments applied at a single phytomer on axillary bud outgrowth in Trifolium repens L.

We studied the effects of light quality and defoliation on the rate of phytomer appearance and axillary bud outgrowth in white clover. The treatments were applied to one phytomer, a phytomer being defined as the structural unit comprising a node, internode, axillary bud, subtending leaf and two nodal root primordia. Light of a low red:far-red (R:FR) ratio (0.27) was applied to a target phytomer either (i) within the apical bud and then to the axillary bud after emergence of the phytomer from the apical bud, or (ii) to the axillary bud only after emergence. The light conditions were directed to these specific parts of the plant by collimating light from small FR light-emitting diodes; with this technique we were able to change the light quality without any change in the level of photosynthetically active radiation. The subtending leaf of the target phytomer was retained or defoliated when it had emerged from the apical bud. FR light applied from the time the phytomer was within the apical bud caused a delay in branch appearance at the target phytomer. In contrast, direct treatment of the axillary bud with FR light after it had emerged from the apical bud did not result in any delay in branch appearance. As the light treatment of the apical bud may have changed the light environment of any of the organs contained in the bud we were unable to ascribe the delay in branch appearance to light perception by any particular organ. However, indirect evidence leads to the conclusion that the likely site of light perception was the developing leaf subtending the axillary bud while it was the outermost phytomer within the apical bud. These results do not support the hypothesis that the R:FR ratio of light incident at an axillary bud site is the environmental factor that controls bud development. Defoliation of the unfolding leaf reduced the rate of phytomer appearance on the main stolon but had no immediate effect on branch appearance. As a consequence there was a reduction in the number of phytomers between the stolon apical meristem and the first phytomer with a branch. This is frequently taken to indicate a relaxation of apical dominance, but in this case was found not to involve a direct effect on bud activity. A current model of white clover growth suggests that there is integration of activity between apical meristems but independence of activity and response to the local micro-environment by axillary buds. In contrast, we found that (i) defoliation reduced phytomer appearance only at the main stolon apical meristem and not at all the meristems in the plant and (ii) that a change in the local light environment of an axillary bud had no discernible effect on bud activity once the bud had emerged from the apical bud but could delay branching if applied before emergence. These results are at variance with the predictions of the model.     

Root growth, developmental changes in the apex, and hydraulic conductivity for Opuntia ficus-indica during drought

Developmental changes in the root apex and accompanying changes in lateral root growth and root hydraulic conductivity were examined for Opuntia ficus-indica (L.) Miller during rapid drying, as occurs for roots near the soil surface, and more gradual drying, as occurs in deeper soil layers. During 7 d of rapid drying (in containers with a 3-cm depth of vermiculite), the rate of root growth decreased sharply and most root apices died; such a determinate pattern of root growth was not due to meristem exhaustion but rather to meristem mortality after 3 d of drying. The length of the meristem, the duration of the cell division cycle, and the length of the elongation zone were unchanged during rapid drying. During 14 d of gradual drying (in containers with a 6-cm depth of vermiculite), root mortality was relatively low; the length of the elongation zone decreased by 70%, the number of meristematic cells decreased 30%, and the duration of the cell cycle increased by 36%. Root hydraulic conductivity ( L _P) decreased to one half during both drying treatments; L _P was restored by 2 d of rewetting owing to the emergence of lateral roots following rapid drying and to renewed apical elongation following gradual drying. Thus, in response to drought, the apical meristems of roots of O. ficus-indica near the surface die, whereas deeper in the substrate cell division and elongation in root apices continue. Water uptake in response to rainfall in the field can be enhanced by lateral root proliferation near the soil surface and additionally by resumption of apical growth for deeper roots.     

Relationships between root diameter,root length and root branching along lateral roots in adult,field-grown maize

Background and Aims Root diameter, especially apical diameter, plays an important role in root development and function. The variation in diameter between roots, and along roots, affects root structure and thus the root system’s overall foraging performance. However, the effect of diameter variation on root elongation, branching and topological connections has not been examined systematically in a population of high-order roots, nor along the roots, especially for mature plants grown in the field.Methods A method combining both excavation and analysis was applied to extract and quantify root architectural traits of adult, field-grown maize plants. The relationships between root diameter and other root architectural characteristics are analysed for two maize cultivars.Key Results The basal diameter of the lateral roots (orders 1–3) was highly variable. Basal diameter was partly determined by the diameter of the bearing segment. Basal diameter defined a potential root length, but the lengths of most roots fell far short of this. This was explained partly by differences in the pattern of diameter change along roots. Diameter tended to decrease along most roots, with the steepness of the gradient of decrease depending on basal diameter. The longest roots were those that maintained (or sometimes increased) their diameters during elongation. The branching density (cm^–1) of laterals was also determined by the diameter of the bearing segment. However, the location of this bearing segment along the mother root was also involved – intermediate positions were associated with higher densities of laterals.Conclusions The method used here allows us to obtain very detailed records of the geometry and topology of a complex root system. Basal diameter and the pattern of diameter change along a root were associated with its final length. These relationships are especially useful in simulations of root elongation and branching in source–sink models.     

Types, structure and potential for axial water flow in the deepest roots of field-grown cereals

Deep root systems that extend into moist soil can significantly increase plant productivity. Here, the components of soil-grown root systems of wheat, barley and triticale are characterized, and types and water conducting potential of deep roots in the field are assessed. Root system components were characterized in plants grown in soil in PVC tubes, based on their origin and number and the arrangement of xylem tracheary elements (XTE) viewed using fluorescence microscopy. A new nomenclature is proposed. Deep roots were harvested in the field, and root types of the current crop and remnant roots from previous crops were identified by fluorescence and cryo-scanning electron microscopy. Four types of axile (framework) and five types of branch root were distinguished in the three cereals. Six per cent of deep roots were axile roots that originated from the base of the embryo; 94% were branch roots, of which 48% had only two XTE (10 microm diameter), and thus potentially low axial flow. Only 30% of roots in the cores were from the current crop, the remainder being remnants. Selection for more deep-penetrating axile roots and increased vascular capacity of deep branches is of potential benefit. Conventional root-length density measurements should be interpreted and applied cautiously.     

Root apical diameter and root elongation rate of rubber seedlings (Hevea brasiliensis) show parallel responses to photoassimilate availability

The hypothesis that root apical diameter may be used to evaluate root growth potential was tested. Temporal variations in the apical diameter of individual roots of rubber seedlings ( Hevea brasiliensis ) were studied together with their elongation patterns, using root observation boxes under controlled conditions. This study confirmed the overall positive correlation between apical diameter and growth rale. Moreover, the two parameters, varied in the same way during the life of a given root. For roots with short growth duration, there was a parallel quick decrease in both apical diameter and elongation rate, whereas roots that grew for longer periods showed synchronous fluctuations for both parameters. Since the mean values for the secondary roots within a root system exhibited the same trends, variations in apical diameter and elongation rates should depend on factors influencing the whole root system. When related to shoot rhythmic growth, both apical diameter and elongation rates were depressed during the periods of leaf growth. These effects were enhanced and/or prolonged by shading, hence reinforcing the hypothesis that this development depends on assimilate availability. Such results can be interpreted in terms of a source-sink relationship within the whole plant by considering the apical diameter, representing the size of the meristem related to the number of rneristematic cells, as an indicator of each root's growth potential.     

Factors affecting the establishment and maintenance of ‘Titan’ red raspberry root organ cultures

Root organ cultures of red raspberry (Rubus ideaus cv. Titan) were established from two cm long terminal segments of adventitious roots induced in vitro on micropropagated shoot cultures. Growth of root organ cultures was almost entirely by initiation and elongation of lateral roots rather than by terminal elongation from the apical meristem of the original root explants. Indolebutyric acid (IBA) was required for lateral root initiation and elongation. The optimal IBA concentration for lateral root initiation decreased from 0.5 to 0.1 mg/L from the first to the fifth passage in culture. Two cm root explants initiated more lateral roots per unit length than four cm explants. Liquid Anderson's medium was superior to other basal nutrient and vitamin formulations tested. Root growth in liquid culture was stimulated by aeration. Root organ cultures also grew on media solidified with gelrite or agar. A rapid transfer technique was developed for subculturing these which involved cutting and transferring 1 cm discs of roots and underlying medium. Adventitious bud formation occurred spontaneously, but sporadically in liquid cultures, and was not influenced by cytokinin, auxin (spat) light, or chilling.     

Roots of willow (Salix viminalis L.) show marked tolerance to oxygen shortage in flooded soils and in solution culture

Responses to soil flooding and oxygen shortage were studied in field, glasshouse and controlled environment conditions. Established stools ofSalix viminalis L., were compared at five field sites in close proximity but with contrasting water table levels and flooding intensities during the preceding winter. There was no marked effect of site on shoot extension rate, time to half maximum length or final length attained. When rooted cuttings were waterlogged for 4 weeks in a glasshouse, soil redox potentials quickly decreased to below zero. Shoot extension was slowed after a delay of 20 d, while, in the upper 100 mm of soil, formation and outgrowth of unbranched adventitious roots with enhanced aerenchyma development was promoted after 7 d. At depths of 100–200 mm and 200–300 mm, extension by existing root axes was halted by soil flooding, while adventitious roots from above failed to penetrate these deeper zones. After 4 weeks waterlogging, all arrested root tips recommenced elongation when the soil was drained; their extension rates exceeding those of roots that were well-drained throughout. Growth in fresh mass was also stimulated. The additional aerenchyma found in adventitious roots in the upper 100 mm of soil may have been ethylene regulated since gas space development was inhibited by silver nitrate, an ethylene action inhibitor. The effectiveness of aerenchyma was tested by blocking the entry of atmospheric oxygen into plants with lanolin applied to lenticels of woody shoots of plants grown in solution culture. Root extension was halved, while shoot growth remained unaffected. H Lambers Section editor     

The involvement of the root apex and cytokinins in the control of lateral root emergence in lettuce seedlings

Removal of the apical 3 mm of the primary root of hydroponically-grown lettuce seedlings 3 or 5 days after sowing, prevented further elongation of the root and increased both the number and total length of lateral roots. The length of the lateral zone, i.e. the distance from the base of the parent root to the lateral nearest the tip, except on one occasion, remained the same as the control in both 3 and 5 day treatments, until the length of the decapitated root (which had ceased elongating) became limiting.Zeatin applied via the roots, at a concentration range from 3 × 10^–10 M to 10^–8 M reduced tap root extension growth at all concentrations. Lateral root emergence was enhanced by low zeatin concentrations and retarded by higher ones. In general, the lateral zone length was the same in cytokinin-treated plants as in untreated controls.     

The formation,morphology and anatomy of cluster root of Lupinus albus L. as dependent on soil type and phosphorus supply

The effect of phosphorus supply on the formation, morphology and anatomy of cluster roots of Lupinus albus L. cv Ultra grown in a loam and two sandy soils was examined relative to its effect on total root length, shoot weight and the phosphorus concentration of the shoots. The loam soil was most conducive to the formation of cluster roots. Cluster roots growing in the sandy soils developed to a lesser extent on plants of an equivalent phosphorus status, suggesting that some biotic or abiotic factors independent of phosphorus supply were also operating. The presence of mature cluster rootlets on a length of lateral root increased the root surface area by 14–22 times of an equal length of lateral roots not bearing cluster rootlets. The application of phosphorus decreased cluster-root length, whereas total root length showed a steady increase. There was an inverse relationship between cluster-root production and phosphorus concentration in shoots ranging from 2 to 8.5 mg g^–1 with the critical phosphorus level for maximum shoot growth being around 2.5 mg g^–1. Cluster roots formed in solution culture were not well developed in comparison with those grown in the loam soil or nutrient solution with added loam soil. The organisation of the cluster rootlet was similar to that of the lateral roots. Mature rootlets lacked an apical meristem and a vascular cambium with a reduced root cap and cortical tissue.     

Relationships among shoot sinks for resources exported from nodal roots regulate branch development of distal non-rooted portions of Trifolium repens L

Two manipulative experiments tested hypotheses pertaining to the correlative control exerted by nodal roots on branch development of the distal non-rooted portion of Trifolium repens growing clonally under near-optimal conditions. The two experiments, differing in their pattern of excision to manipulate the number of branches formed at the first 9-10 phytomers distal to the youngest nodal root, each found that after 20 phytomers of growth the total number of lateral branches formed on the primary stolon remained between five and seven regardless of where the branches formed along the stolon. Additional treatments established that nodal roots influenced branch development via relationships among shoot sinks for the root-supplied resources rather than through variation in the supply of such resources induced by fluctuations in photosynthate supply to roots from branches. Regression analysis of data pooled from treatments of both experiments confirmed that shoot-sink relationships for root- supplied resources controlled the branching processes on the non-rooted portion of plants. A disbudding treatment, which removed all the apical and axillary buds present on basal branches, but left other branch tissues intact, increased branch development of the apical region in the same way as did complete excision of the basal lateral branches. The apical buds and the elongation processes occurring immediately proximal to the buds were thus identified as strong sinks for the root-supplied resources. Such results suggest that branch development on the non-rooted shoot portion distal to the youngest nodal root is regulated by competition among sinks for root-derived resources, of limited availability, necessary for the processes of elongation of axillary buds and the primary stolon apical bud.     

Architectural analysis and modelling of the branching process of the young oil-palm root system

An architectural analysis of the root system of young oil-palm (Elaeis guineensis Jacq.) seedlings was made. In this analysis, root branching was modelled by a Markov chain (discrete-time, discrete-state space stochastic process). This study has been realized on radicles of young oil-palm seedlings which were considered as main axes which branch. We defined an elementary length unit as the smallest length between two successive lateral roots. The model was based on the analysis of a sequence of events, each event being indexed by the rank of the elementary length unit on the main axis. An event was defined as the state of the length unit, chosen between unbranched state and three branched-state categories. The branching process of the oil-palm radicle was modelled by a four-state first-order Markov chain. Consequently, the state of an elementary length unit depended only on the state of the previous one. The Markov chain was homogeneous, i.e. the transition probabilities did not depend on the rank of the elementary length unit.This study allowed us to identify a probabilistic model of root branching which was the first step in the elaboration of a stochastic model of the architecture of the oil-palm root system.     

玉米初生根向水性诱导优化试验研究

为了研究湿度梯度对根系向水性反应的影响,采用Takahashi and Scott于1993年创建的方法,设置以下3个试验:1)向水性诱导物不同倾斜角试验;2)根系距向水性诱导物不同距离试验;3)根尖距底部饱和K2CO3溶液不同距离试验。同时,还研究了根长和根系延伸速率对根系向水性弯曲的影响。结果表明,用饱和K2CO3溶液控制湿度时根系的向水性弯曲度明显大于纯水。随着诱导物倾斜角的增大,向水性弯曲增强。与距诱导物3 mm和6 mm相比,根系直接接触诱导物时表现出最大的向水性反应。与根尖距底部盐溶液6 cm相比,相距4 cm时向水性弯曲度增大,这些与根尖周围的湿度梯度增大有关。当根长为1.0、1.5、2.0、2.5、3.0 cm时,短根比长根表现出更大的向水性反应,这可能与其较慢的延伸速率为根系对湿度梯度的反应提供了更充足的时间有关。为了验证这个假说,用相同长度的根系、通过控制不同温度进行试验,结果表明根系的向水性弯曲随温度升高而降低。可见,玉米初生根的向水性反应受环境和根系发育阶段两方面影响。当根系相距诱导物较近、根系周围的湿度梯度较大时,根系向水性反应更强。而且,具有较小延伸速率根系的向水性反应更大。考虑到干旱条件下根系伸长慢、且土壤中湿度梯度大,因而可以认为干旱条件下根系的向水性生长在玉米吸收水分中有重要作用。同时,对根系向水性诱导方法的优化有助于其生理机制的进一步研究。