Asynchronicity in root and shoot phenology in grasses and woody plants

Phenology is central to understanding vegetation response to climate change, as well as vegetation effects on plant resources, but most temporal production data is based on shoots, especially those of trees. In contrast, most production in temperate and colder regions is belowground, and is frequently dominated by grasses. We report root and shoot phenology in 7‐year old monocultures of 10 dominant species (five woody species, five grasses) in southern Canada. Woody shoot production was greatest about 8 weeks before the peak of root production, whereas grass shoot maxima preceded root maxima by 2–4 weeks. Over the growing season, woody root, and grass root and shoot production increased significantly with soil temperature. In contrast, the timing of woody shoot production was not related to soil temperature (r=0.01). The duration of root production was significantly greater than that of shoot production (grasses: 22%, woody species: 54%). Woody species produced cooler and moister soils than grasses, but growth forms did not affect seasonal patterns of soil conditions. Although woody shoots are the current benchmark for phenology studies, the other three components examined here (woody plant roots, grass shoots and roots) differed greatly in peak production time, as well as production duration. These results highlight that shoot and root phenology is not coincident, and further, that major plant growth forms differ in their timing of above‐ and belowground production. Thus, considering total plant phenology instead of only tree shoot phenology should provide a better understanding of ecosystem response to climate change.     

Root development and configuration in intensively managed radiata pine plantations

Summary In south-east Australia, where radiata pine (Pinus radiata D. Don) is grown on sandy soils low in nutrients and short of water, early establishment, and rapid growth to canopy closure lead to increased productivity. At this stage demands for nutrients and water are high, and trees respond vigorously to silvicultural inputs.For several months after transplanting in winter roots are confined within a narrow planting wedge, low temperature restricts new root growth and slows recovery from water stress in plants. From spring, depending upon the configuration and vigour of the roots transplanted, lateral roots extend radially throughout the soil.Although there were small decreases in concentration of roots radially from the stems of very young trees, such spatial differences disappeared between ages 2 and 3, so that rooting density was independent of distance from the stem. The pattern of vertical distribution of lateral roots was not influenced by age and 80–90% of the lateral roots were within the top 30 cm soil. Roots developed rapidly as the trees grew towards canopy closure, but in general the rooting densities of these pines are among the lowest reported for plants. In rapidly growing trees approaching canopy closure, the secondary thickening of the lateral roots was sufficient to double the weight of roots without altering root length.Knowledge about root growth and root configuration during the early phase of plantation development will assist management decisions where intensive silviculture is practiced, and hence ensure the most efficient use of nutrients and water.     

Tree and grass rooting patterns in an African humid savanna

Abstract. Spatial and temporal soil partitioning between roots of the two savanna plant components, i.e. trees and grasses, were investigated in a West African humid savanna. Vertical root phytomass distribution was described for grass roots, large (> 2 mm) and fine (< 2 mm) tree roots, in open sites and beneath tree canopies. These profiles were established monthly over one year of vegetation growth. Natural ¹³C abundance measurement was used to determine the woody/herbaceous phytomass ratio in root samples. Tree and grass root distributions widely overlapped and both were mostly located in the top 20 cm of the soil. Grass root phytomass decreased with depth whereas woody root phytomass peaked at about 10 cm depth. No time partitioning was detected. These structural results do not support the hypothesis of soil resource partitioning between trees and grasses and are thus consistent with functional results previously reported.     

Growth and morphology of eucalypt seedling-roots, in relation to soil strength arising from compaction

Information on the response of root growth and morphology to soil strength is useful for testing suitability of existing and new tillage methods and/or for selecting plants suitable for a specific site with or without tillage. Although there is extensive published information on the root growth-soil strength relationships for annual agricultural plants, such information is scarce for woody, perennial tree species. The purpose of this study is to examine growth and morphology of the root systems of 17-day-old eucalypt seedlings with respect to variation in soil strength. Soil strength in this study was varied by compaction of a well-aggregated clay soil to bulk densities of 0.7–1.0 Mg m^-3 whilst maintaining adequate water availability and aeration for plant growth. Lengths and tip-diameters of primary and lateral roots were measured on the excavated root systems of seedlings.With increase in bulk density and also soil strength (expressed as penetrometer resistance), total length of primary and lateral roots decreased. There were 71 and 31% reduction in the lengths of primary and lateral roots respectively with an increase in penetrometer resistance from 0.4 to 4.2 MPa. This indicated primary roots to be more sensitive to high soil strength than the lateral roots. Average length of lateral roots and diameters of both primary and lateral root tips increased with an increase in soil strength as well. There was greater abundance of lateral roots (no. of lateral roots per unit length of primary root) and root hairs with increased soil strength. The observed root behaviour to variable soil strength is discussed in the context of compensatory growth of roots and overall growth of plants.     

Root proliferation characteristics of seven perennial arid-land grasses in nutrient-enriched microsites

We compared root proliferation in fertilized microsites among seven cultivars of five commonly planted cool-desert perennial grass species that differ in productivity and competitive ability. In a greenhouse experiment on nutrient-limited plants, one soil microsite in each pot received distilled water (control) and a second microsite received a rich, complete nutrient solution (fertilized). Roots in and adjacent to the microsites were mapped on Mylar windows for 22 days after the injections to determine the magnitude and timing of response in root length relative growth rates (RGRs). Because we provided adequate water, used a high level of fertilization in the treatment microsites, and conducted the experiments during rapid vegetative growth, the results provide a measure of the relative capacities and maximal rates of the grasses responses to enriched microsites. Root samples were harvested from control and fertilized microsites at the end of the experiment to determine the morphological basis of the proliferation response. In all seven grasses fine roots proliferated in the fertilized microsites faster than in the control microsites. The grasses did not differ in the timing of their response which showed a peak 7–8 days after injection. Although one species, Pseudoroegneria spicata cv. Goldar, had higher maximum root length RGR and higher RGR ratio (RGR in fertilized to RGR in control microsites) 7–8 days after injection, the seven grasses did not differ significantly in the magnitude of root length RGR response to fertilizer integrated over the 22 day experiment. The grasses also did not differ significantly in root morphological changes in fertilized mocrosites. Compared to roots in control microsites, roots in fertilized microsites had greater specific root length, length of secondary roots per length of main axis, number of lateral and sublateral roots per length of main axis, and mean lateral root length. Root proliferation was mainly the result of increased lateral branching and lateral root growth in all seven grasses. The consistency of root proliferation responses among these seven cultivars suggests that differences in the capacity for, maximum rate, or morphological basis of root proliferation are not directly related to ecological characteristics such as productivity and competitive ability. Other aspects of root response to nutrient enrichment, such as differential responses as a function of microsite nutrient concentration, plant phenology, plant nutrient status, or specific nutrient element(s), may still be important, but further experiments are required to determine whether different responses to enriched soil microsites among species correspond with know species differences in ecological characteristics.     

Water deficit accelerates determinate developmental program of the primary root and does not affect lateral root initiation in a Sonoran Desert cactus (Pachycereus pringlei, Cactaceae)

Determinate root growth is an important adaptation feature for seedling establishment in some Cactaceae. We show that seedlings of Pachycereus pringlei have primary roots with a stable determinate developmental program. How water stress affects determinate root growth and lateral root development has not been studied. Here we address this question. Root growth was analyzed in plants growing in vitro under well-watered and water-deficient (created by polyethylene glycol) growth conditions. Under severe water stress roots terminated their growth earlier and the rate of growth was significantly decreased as a result of inhibition of both cell elongation and cell production. Under severe water stress the number of lateral roots and primordia per millimeter of primary root was 1.5-1.7 times greater than under well-watered conditions; however, the total number of lateral roots and primordia was the same under all conditions. Lateral roots resembled root spurs found in some Opuntioideae. Analysis of the dynamics of meristem exhaustion indicated that initial-cell activities are required for the maintenance of proliferation before meristem exhaustion. We conclude that lateral root formation is a stable developmental process resistant to severe water stress and that water stress accelerates the determinate developmental program of the primary root. Both of these features appear to be important for successful seedling establishment in a desert.     

Increased nutrient supply facilitates acclimation to high-water level in the marsh plant Deyeuxia angustifolia: The response of root morphology

Both water level and nutrient availability are important factors influencing the growth of wetland plants. Increased nutrient supply might counteract the negative effects of flooding on the growth of the fast-growing species. Experimental evidence is scarce and the mechanism is far from clear. The aim of this study is to identify the role of nutrient availability in acclimation to high-water level by investigating the growth and root morphology of the marsh plant Deyeuxia angustifolia, one of the dominant species in the Sanjiang Plain, China. Experimental treatments included two water levels (0 and 10 cm, relative to soil surface) and three levels of nutrient supply (0, 0.5 and 1 g fertilizer per container). High-water level usually led to decreased biomass accumulation, shoot mass and root mass, whereas biomass accumulation was unaffected by water level at the highest nutrient level, indicating that high-nutrient availability played a role in compensating for the growth loss induced by the high-water level. Increased nutrient supply led to decreased root length in 0 cm water-level treatments, but root length increased with nutrient supply in the 10 cm water-level treatments. High-water level usually led to a lower lateral root density, lateral root:main root length ratio and the diameter of main roots and laterals, whereas increased nutrient supply resulted in thicker main roots or laterals, and a higher total root length, lateral root density and lateral root:main root length ratio. These data indicate that the growth of D. angustifolia is restrained by high-water level, and that increased nutrient supply not only ameliorates root characteristics to acclimate to high-water level but also results in a high-total root length to facilitate nutrient acquisition.     

Effect of Phosphorus Deficiency Stress on Rice Lateral Root Growth and Nutrient Absorption

Six rice (Oryza sativa L.) genotypes with different performances under phosphorus (P) deficiency stress were tested in mixed growth medium of vermiculite and sand under different conditions of P supply to evaluate the effects of P deficiency stress on lateral root growth and the relations between lateral root growth induced by P deficiency and P absorption. The results showed that elongation and development of lateral root were induced by P deficiency. There was significant genotypic variation in lateral root growth in response to P deficiency. A significant positive correlation was observed between the increase of lateral root length per cm of nodal root and the increase of root surface area per cm of nodal root (RSAP), while no significant correlation was observed between the increase of lateral root number per cm of nodal root and the increase of RSAP. The result suggested that the increase of root surface area under P deficiency condition could be mainly attributable to the increase of lateral root length induced. P uptake was significantly positively correlated with the total root surface area and positively correlated with the total lateral root length and the total lateral root number under P deficiency, which implied that elongation and development of lateral root were important to the ability of P uptake from growth medium where P supply was poor. Analysis of soluble sugar content indicated that P deficiency stress changed the distribution of carbohydrate between roots and shoots.     

Growth and root responses of woody species to rocky substrate: implications for gully rehabilitation

Gullies formed in the Velhas River basin in Brazil have been filled with urban construction waste for physical stabilisation purposes. Aimed at rehabilitating gullies, we selected woody species from the Brazilian Cerrado that can grow on rocky substrates under greenhouse conditions. An assessment was made regarding plant growth in both rocky and natural soil substrates by analysing the height, diameter, fresh and dry weights of shoots and roots, plant water content, root occupation and architecture. Principal component analysis and Chi-squared tests segregated rock-tolerant species based on the specific influence on root dry and fresh weights. Fast-growing species reduced the emergence of their lateral roots under rocks, compromising their growth in height and biomass production. In contrast, slow-growing woody species were particularly suitable for gully rehabilitation because these species exhibited a genetic pattern of low lateral root emergence that prevented damage to their roots. Most slow-growing species demonstrated a similar growth pattern in both substrates, and some of them, such as Copaifera langsdorffii, achieved better growth in height and biomass production on rocks than on soil, a finding attributed to the root plastic response involving primary root elongation and lateral root emergence. Therefore, slow-growing species are recommended for gully rehabilitation procedures.     

Variations in maturation strains and root shape in root systems of Maritime pine (Pinus pinaster Ait.)

 In order to determine if different types of wood were being laid down in the root system of Maritime pine (Pinus pinaster Ait), in response to wind loading, longitudinal residual maturation strains (LRMS), indicating the existence of mechanical stress in developing wood cells, were measured in the trunk and lateral roots. Two age groups of trees (5- and 13-year- old) were compared. LRMS were greater in the trunk and roots of 13-year-old trees than in 5-year-old trees. This phenomenon may be due to increased competition between older trees. LRMS in leeward roots of both age-groups were positive i.e. the wood cells had developed under compression, as also occurs in reaction wood of gymnosperms. As leeward roots are placed under compression during tree sway, an abnormal type of wood may form in the roots in order to counteract the increased stress. In other roots, the strains were negative i.e. the cells had developed under tension, as occurs in normal wood. In the roots of younger trees, LRMS were also positive nearer the stem, thus indicating that wood formation may also be influenced by bending stresses experienced in this zone. In addition to LRMS measurements, radial growth in roots was examined in order to determine the influence of mechanical loading on secondary growth. In older trees, there was a significant increase of 34% in woody growth below the biological centre, compared to that above. This eccentricity is unlike that found in most other tree species, where secondary growth is usually greater on the upper side of the root. However, Maritime pine has a tap root, which will alter the pattern of stress within the root system. Under wind loading, a concentration of mechanical stress will develop at the bases of the stem, lateral roots and tap root. Received: 7 July 1997 / Accepted: 11 December 1997     

Citrus root responses to localized drying soil: A new approach to studying mycorrhizal effects on the roots of mature trees

Because fine roots tend to be concentrated at the soil surface, exposure to dry surface soil can have a large influence on patterns of root growth, death and respiration. We studied the effects of arbuscular mycorrhizas (AM) formation on specific root length (SRL), respiration and mortality of fine roots of bearing red grapefruit (Citrus paradisi Macf.) trees on Volkamer lemon (C. volkameriana Tan. & Pasq.) rootstock exposed to drying soil. For each tree, the fine roots were removed from two woody lateral roots, the roots were surface sterilized and then each woody root was placed in a separate pair of vertically divided and independently irrigated soil compartments. The two split-pot systems were filled with sterilized soil and one was inoculated with arbuscular mycorrhizal fungi (Glomus etunicatum/G. intraradices). New fine lateral roots that emerged from the woody laterals were permitted to grow inside the pots over a 10-month period. Irrigation was then removed from the top compartment for a 15-week period. At the end of the study, roots inoculated with AM fungi exhibited about 20% incidence of AM formation, whereas the uninoculated roots were completely void of AM fungi. Arbuscular mycorrhizal roots exhibited lower SRL, lower root/soil respiration and about 10% lower fine root mortality than nonmycorrhizal roots after 15 weeks of exposure to dry surface soil. This study demonstrates the feasibility of examining mycorrhizal effects on the fine roots of adult trees in the field using simple inexpensive methods.     

Influence of wind loading on root system development and architecture in oak (Quercus robur L.) seedlings.

The effect of wind loading on seedlings of English oak (Quercus robur L.) was investigated. Instead of using a traditional wind tunnel, an innovative ventilation system was designed. This device was set up in the field and composed of a rotating arm supporting an electrical fan, which emitted an air current similar to that of wind loading. Oaks were sown from seed in a circle around the device. A block of control plants was situated nearby, and was not subjected to artificial wind loading. After 7 months, 16 plants from each treatment were excavated, and root architecture and morphological characteristics measured using a 3D digitiser. The resulting geometrical and topological data were then analysed using AMAPmod software. Results showed that total lateral root number and length in wind stressed plants were over two times greater than that in control trees. However, total lateral root volume did not differ significantly between treatments. In comparing lateral root characters between the two populations, it was found that mean root length, diameter and volume were similar between the two treatments. In trees subjected to wind loading, an accentuated asymmetry of root distribution and mean root length was found between the windward and leeward sides of the root system, with windward roots being significantly more numerous and longer than leeward roots. However, no differences were found when the two sectors perpendicular to the wind direction were compared. Mean tap root length was significantly higher in control samples compared to wind stressed plants, whilst mean diameter was greater in the latter. Wind loading appears to result in increased growth of lateral roots at the expense of the tap root. Development of the lateral root system may therefore ensure better anchorage of young trees subjected to wind loading under certain conditions.     

不同刈割强度对牧草地上部和地下部生长性状的影响

研究了不同刈割强度对牧草地上部和地下部生长状况的影响.结果表明,适度刈割可提高牧草地上部植株的再生能力.刈割后牧草再生叶片的叶绿素总量变化不大,而叶绿素a/b比值有所增加,轻刈割和重刈割的牧草叶绿素a/b比值分别增至1.59∶1和1.52∶1、不刈割为1.22∶1,有利于增强植物的光合作用.与不刈割处理相比,在刈割初期,重刈割处理下柱花草根系总长、总表面积和平均直径分别下降了54.9%、66.5%和27.2%,根系活力显著下降;但在中后期,刈割处理的牧草地下部根系形态指标活力可恢复到更高的水平.从一年两次收获的累计生物量来看,以轻刈割最高,为3 179.8 g·m^-2,重刈割次之,为3 006.1 g·m^-2,不刈割最低,为2 936.98 g·m^-2,说明一年两次刈割可以提高牧草产量.     

Growth rates,biomass and distribution of selected woody plant roots in Burkea africana-Ochna pulchra savanna

Woody plants in an African Burkea africana-Ochna pulchra savanna on deep sandy soil were found to have characteristically bimorphic root systems. The shallow lateral root component was often well developed and roots extended up to seven times the extent of the plant canopy in several species. Exponential tapering of lateral roots was found in Terminalia sericea. The wide-ranging roots, together with the high degree of multispecies root system interpenetration, result in the so-called, open grassy areas in the savanna mosaic often containing a competitively significant woody plant component. Root systems of Ochna pulchra were found to be relatively specialized and included: negatively geotropic, superficial roots; sinker roots to bedrock; high suckering response to damage in roots; belowground lignotuber-type organs; and sustained subterranean interconnections between some aboveground stems. These features are likely to contribute substantially to the resilience of this plant species to various climatic and veld management stress factors. Root/shoot mass ratios averaged unity but depended on plant size and aboveground growth form in Ochna pulchra. The dependence of these ratios on sizes of plant also applied to plant clones. Initiation of root tip growth occurred in early summer in one year and late spring in another. Main root tip growth occurred in late summer and early autumn, well after completion of most growth of leafy shoots in spring. It is suggested that some active uptake of water and nutrients by non-extending roots allows this form of phased growth in the plant. In an analysis of the seasonal growth of individual root tip systems, it was clear that transitory states of rest occur in fine root development but that these are far more frequent in the branching (and hence proliferation) of roots than in the continuing development of any root axis.Nomenclature follows the present system of the Botanical Research Institute, Pretoria, and the Flora of Southern Africa.I thank M.D. Panagos, P.S. Carr and J. Steyn for assistance at various stages of this work.     

Influence of rocks on soil temperature,soil water potential,and rooting patterns for desert succulents

Summary At a site in the Sonoran Desert, subterranean rocks and exposed boulders affected soil water potential as well as root morphology and distribution. For Agave deserti, the number of lateral roots per unit length of main root was 11 times higher under rocks and six times higher alongside rocks than in rock-free regions. Total root length per unit soil volume for Echinocereus engelmannii averaged 3-fold higher within 1 cm of boulders than 5 cm away, where the soil was drier. The total length of lateral roots per unit length of main root for Ferocactus acanthodes was 4.2 m m^–1 under rocks but only 0.8 m m^–1 in rock-free regions. The number of lateral roots per unit length of main root for Opuntia acanthocarpa was 7-fold higher alongside rocks than in rock-free regions and even higher under rocks. For transplanted and watered A. deserti, the number of new main roots produced per 1–2 month interval averaged 13 for five plants on the north side of boulders, 8 on the south side, 11 for five plants with half of their roots under rocks, 2 for those with half of their roots over rocks, and 3 for the control plants without rocks. Laboratory experiments showed that the soil water potential under rocks for 10 and 30 mm waterings stayed above –0.5 MPa for 13 and 19 d longer, respectively, than for regions away from rocks. The shortwave absorptance of granitic rocks from the field site was 0.82, the thermal conductivity coefficient was 1.50 W m^–1 °C^–1, and the volumetric heat capacity was 1.75 MJ m^–3 °C^–1. Field measurements indicated that 5-cm-thick buried rocks decreased the diel variation in soil temperatures on their undersurface by only 0.4° C compared with soil. Thus, the primary influence of rocks at the field site on root proliferation and branching for the four species was apparently caused by influences on soil water content.     

低磷胁迫对水稻苗期侧根生长及养分吸收的影响

用蛭石与石英砂作为混合培养介质研究了低磷胁迫对水稻（Oryza sativa L.)苗期侧根发生发育的影响及其与磷吸收的相关关系。结果表明：低磷对水稻的侧根发生发育具有明显的诱导作用及基因型差异。相关性分析表明：单位侧根长度的增加与单位根表面积的增大极显相关,而单位侧根数量的增多与单位根表面积的增大无显的相关性。表明单位根表面积的增加主要来自于单位侧根的伸长。侧根参数与磷含量的相关性分析表明：低磷条件下,侧根总长度和侧根数量都与植株磷含量存在显的正相关,根系总表面积与磷含量存在极显的正相关。表明在低磷条件下,侧根的发生发育对水稻的磷吸收具有重要的作用。根系和地上部的可溶性糖含量分析表明;低磷胁迫改变了同化物在地上部和根系的分配。生物量测定表明：低磷胁迫显增大了植株的根冠比。     

Genotypic Variation in Root Growth Angle in Rice (Oryza sativa L.) and its Association with Deep Root Development in Upland Fields with Different Water Regimes

Deep root development, which is important for the drought resistance in rice (Oryza sativa L.), is a complex trait combining various root morphologies. The objective of this study was to elucidate genotypic variation in deep root development in relation to morphological indicators such as vertical root distribution and root growth angle. Two experiments were conducted: one on upland fields, and one in pots and fields. In experiment 1, the root systems of six rice cultivars on upland fields were physio-morphologically analyzed under different water regimes (irrigated and intermittent drought conditions during panicle development). In experiment 2, cultivar differences in root growth angles were evaluated with 12 cultivars using the basket method under irrigated conditions. No cultivar × environment interactions were found for total root length or deep root length between irrigated and drought conditions in experiment 1. This suggests that constitutive root growth, which is genetically determined, is important for deep root development under intermittent drought conditions during reproductive stage. Among root traits, the deep root ratio (i.e., deep root weight divided by total root weight) was most closely related to deep root length under both water regimes. This suggested that vertical root distribution constitutively affects deep root length. Significant genotypic variation existed in the nodal root diameter and root growth angle of upland rice in experiment 2. It was considered that genotypes with thick roots allocated more assimilates to deep roots through root growth angles higher to the horizontal plane on upland fields. This is the first report on genotypic variation in the root growth angle of rice on upland fields. It should prove useful for rough estimations of genotypic variation in the vertical root distribution of upland rice because root growth angle is rapidly and easily measured.     

Assessment of a rapid method,using soil cores,for estimating the amount and distribution of crop roots in the field

Summary A study was made of the relationship between the number of roots (N_r) observed on unit area of the freshly exposed, horizontal faces of soil cores, and the amounts of roots (per unit volume) present in the same cores. Soil cores, 7 cm diameter, were extracted to depths of 1 m from cereal crops in 1976 at three field sites located on clay soils. Sampling was either at the start of stem elongation, or at anthesis. Estimates of root length per unit soil volume (L) were derived from N_r by assuming random orientation of roots in the soil.Values of L were found to be highly correlated with the measured lengths of both the main roots (root axes) and the total roots (axes and laterals) washed from the soil at a given growth stage, for each of the soils. On average, L was 3.3 times the length of root axes washed from the soil, and was 0.42 times the length of total roots, but there was appreciable variation between different growth stages and field sites. Possible factors giving rise to differences between L and the measured lengths of roots are discussed. Estimates of root length from observation of soil cores may nonetheless provide a suitable basis for rapidly comparing therelative distribution of roots down the soil profile under field conditions.     

Longevity and turnover of roots in the shortgrass steppe: influence of diameter and depth

We used minirhizotrons to determine patterns of root longevity andturnover for the perennial bunchgrass Bouteloua gracilisinthe shortgrass steppe of eastern Colorado, USA. We hypothesized that rootlongevity would be partially controlled by root diameter, following previouslyobserved patterns in woody plants. In addition, we hypothesized that rootturnover would be greatest in surface soil horizons and decrease with depth dueto variation in soil moisture availability and temperature. Root longevity wascorrelated with root diameter. Median life span of roots > 0.4mm was approximately 320 days, while roots < 0.2mmhad a median life span of 180 days. There was approximately a 6%decreasein the likelihood of mortality with a 0.10-mm increase inroot diameter, controlling for the effect of depth in the soil profile. Rootlength production and mortality were highest in the upper20 cm of the soil profile and decreased with depth.However,because root length density also decreased with depth, there were nosignificantdifferences in turnover rate of root length among sampling intervals. Turnoverwas approximately 0.86 yr^–1 based on root length production,while turnover was 0.35 yr^–1 using root length mortality as ameasurement of flux. The imbalance between turnover estimates may be aconsequence of the time the minirhizotrons were in place prior to imaging or mayresult from our lack of over-winter measures of mortality. Our worksuggests that Bouteloua gracilis roots have complex lifehistory strategies, similar to woody species. Some portion of the root systemishighly ephemeral, while slightly larger roots persist much longer. Thesedifferences have implications for belowground carbon and nitrogen cycles in theshortgrass steppe.     

Ongoing modifications to root system architecture of Pinus ponderosa growing on a sloped site revealed by tree-ring analysis

Our knowledge of the root system architecture of trees is still incomplete, especially concerning how biomass partitioning is regulated to achieve an optimal, but often unequal, distribution of resources. In addition, our comprehension of root system architecture development as a result of the adaptation process is limited because most studies lack a temporal approach. To add to our understanding, we excavated 32-year-old Pinus ponderosa trees from a steep, forested site in northern Idaho USA. The root systems were discretized by a low magnetic field digitizer and along with AMAPmod software we examined their root traits (i.e. order category, topology, growth direction length, and volume) in four quadrants: downslope, upslope, windward, and leeward. On one tree, we analyzed tree rings to compare the ages of lateral roots relative to their parental root, and to assess the occurrence of compression wood. We found that, from their onset, first-order lateral roots have similar patterns of ring eccentricity suggesting an innate ability to respond to different mechanical forces; more root system was allocated downslope and to the windward quadrant. In addition, we noted that shallow roots, which all presented compression wood, appear to be the most important component of anchorage. Finally, we observed that lateral roots can change growth direction in response to mechanical forces, as well as produce new lateral roots at any development stage and wherever along their axis. These findings suggest that trees adjust their root spatial deployment in response to environmental conditions, these roots form compression wood to dissipate mechanical forces, and new lateral roots can arise anywhere and at any time on the existing system in apparent response to mechanical forces.