The Wageningen Rhizolab — a facility to study soil-root-shoot-atmosphere interactions in crops

A research facility is described for the integrated study of soil-root-shoot-atmosphere relationships in crops. The Wageningen Rhizolab has been in use since 1990, and consists of two rows, each with eight below-ground compartments aligned along a corridor. A rain shelter automatically covers the experimental area at the start of rainfall. Compartments are 125 cm × 125 cm and 200 cm deep. Each compartment has a separate drip irrigation system. Crop canopy photosynthesis, respiration, and transpiration can be measured simultaneously and continuously on four out of eight compartments at a time. Each compartment can be filled with a selected soil material (repacked soil) and is accessible from the corridor over its full depth. Multiple sensors for measuring soil moisture status, electrical conductivity, temperature, soil respiration, trace gases and oxygen are installed in spatial patterns in accordance with the requirements of the experiments. Sensors are connected to control and data-acquisition devices. Likewise, provisions have been made to sample manually the soil solution and soil atmosphere. Root observation tubes (minirhizotrons) are installed horizontally at depth intervals ranging from 5 cm (upper soil layers) to 25 cm (below 1 m). The facility is at present in use to study growth and development of vegetation (crops) in relation to drought, nutrient status, soil-borne diseases, and underground root competition. One important application is the study of elevated CO₂ concentration and climate change and the way they affect crops and their carbon economy. Growth and development of field grown vegetables and winter cover crops are also evaluated. The common aspect of those studies is to gain a better understanding of crop growth under varying environmental conditions, and to collect datasets that may help to improve mechanistic crop growth simulation models that can address suboptimal growth conditions.     

Root turnover in pasture species: chicory,lucerne, perennial ryegrass and white clover

For pastures, root turnover can have an important influence on nutrient and carbon cycling, and plant performance. Turnover was calculated from mini‐rhizotron observations for chicory (Cichorium intybus), lucerne (Medicago sativa), perennial ryegrass (Lolium perenne) and white clover (Trifolium repens) grown in the Manawatu, New Zealand. The species were combined factorially with four earthworm species treatments and a no‐earthworm control. Split plots compared the effects of not cutting and cutting the shoots at intervals. Observations were made c. 18 days apart for 2.5 years. This article concentrates on differences between plant species in root turnover in the whole soil profile to 40 cm depth. At this scale, earthworm effects were generally small and short lived. For ryegrass and white clover, root length and mass were linearly related (R² = 0.82–0.99). For chicory and lucerne, the relationships were poorer (R² = 0.38–0.77), so for those species length turnover may be a poor indicator of mass turnover. Standing root length, total growth and death generally decreased in the sequence ryegrass > lucerne > chicory = white clover. In length terms, scaled turnover (growth divided by average standing root length) generally followed the sequence lucerne > white clover > perennial ryegrass = chicory. Across species the scaled turnover rate averaged 3.4 per year or 0.9% per day. Cutting shoots reduced standing root length, growth and death, but increased scaled turnover. These results indicate fast and prolonged root turnover. For ryegrass and white clover, at least there is need to reappraise how to measure and model shoot : root ratios, dry matter production and carbon cycling.     

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.     

Photosynthesis and root growth in Spartina alterniflora in relation to root zone aeration

Spartina alterniflora Lois. is a dominant species growing in intermediate and saline marshes of the US Gulf coast and Atlantic coastal marshes. S. alterniflora plants were subjected to a range of soil redox potential (Eh) conditions representing a well aerated to reduced conditions in a rhizotron system under controlled environmental conditions. The low soil Eh resulted in inhibition of root elongation shortly after treatment initiation. Root elongation was reduced as soil Eh approached values below ca. +350 mV. Substantial decrease in root elongation was noted when soil Eh fell below +200 mV. Generally, net photosynthetic rate (PN) decreased as soil Eh was reduced, with substantial reductions in PN found when Eh approached negative values. Average PN was reduced to 87, 64, and 44% of control under +340, +245, and -180 mV treatments, respectively. The reductions in root elongation and PN in response to low soil Eh indicated the adverse effects of low soil Eh on plant functioning and the need for periods of soil aeration that allow plants to resume normal functioning. Thus periods of drainage allowing soil aeration during the growing season appear to be critical to S. alterniflora by providing favorable conditions for root growth and gas exchange with important implications for plant carbon fixation. This revised version was published online in June 2006 with corrections to the Cover Date.     

Root growth dynamics linked to above-ground growth in walnut (Juglans regia)

Background and Aims Examination of plant growth below ground is relatively scant compared with that above ground, and is needed to understand whole-plant responses to the environment. This study examines whether the seasonal timing of fine root growth and the spatial distribution of this growth through the soil profile varies in response to canopy manipulation and soil temperature.Methods Plasticity in the seasonal timing and vertical distribution of root production in response to canopy and soil water manipulation was analysed in field-grown walnut (Juglans regia ‘Chandler’) using minirhizotron techniques.Key Results Root production in walnuts followed a unimodal curve, with one marked flush of root growth starting in mid-May, with a peak in mid-June. Root production declined later in the season, corresponding to increased soil temperature, as well as to the period of major carbohydrate allocation to reproduction. Canopy and soil moisture manipulation did not influence the timing of root production, but did influence the vertical distribution of roots through the soil profile. Water deficit appeared to promote root production in deeper soil layers for mining soil water. Canopy removal appeared to promote shallow root production.Conclusions The findings of this study add to growing evidence that root growth in many ecosystems follows a unimodal curve with one marked flush of root growth in coordination with the initial leaf flush of the season. Root vertical distribution appeared to have greater plasticity than timing of root production in this system, with temperature and/or carbohydrate competition constraining the timing of root growth. Effects on root distribution can have serious impacts on trees, with shallow rooting having negative impacts in years with limited soil water or positive impacts in years with wet springs, and deep rooting having positive impacts on soil water mining from deeper soil layers but negative impacts in years with wet springs.     

Effects of liming on rhizosphere chemistry and growth of fine roots and of shoots of sessile oak (Quercus petraea)

Soil acidification can be detrimental to root growth and nutrient uptake, and liming may alleviate such acidification. In the following study, seedlings of sessile oak (Quercus petraea Liebl. M.) were grown in rhizotrons and subjected to liming (L) or gypsum (G) treatments and compared with the control (C). In order to study and interpret the impact of these calcium rich treatments on fine root development and tree growth, the following parameters were assessed: fine root biomass, fine root length, seedling development (height, diameter, leaves), seedling biomass, nutrient content of roots and seedlings, bulk soil and soil solution chemistry and rhizosphere soil chemistry. The results show that liming increased bulk soil pH, exchangeable Mg, Ca and the Ca/Al molar ratio, and decreased exchangeable Al, mainly in the A-horizon. Gypsum had a similar but smaller impact on exchangeable Al, Ca, H⁺ and the Ca/Al molar ratio in the A-horizon, but reacted with depth, so that exchangeable Mn, Mg and Ca were increased in the B-horizon. In the rhizosphere, the general pattern was determined by the treatment effects of the bulk soil. Most elements were more concentrated in the rhizosphere than in bulk soil, except for Ca which was less concentrated after liming or gypsum application. In the B-horizon rhizosphere pH was increased by the treatments (L > G,C) close to the root tips. Furthermore, the length of the zone with a positive root-induced pH increase was greater for the limed roots as compared with both the other treatments. Fine root growth was stimulated by liming (L > G,C) both in terms of biomass and length, whereas specific root length was not obviously affected apart from the indication of some stimulation after liming at the beginning. The live:dead ratio of fine roots was significantly higher in the limed rhizotrons as compared to the control (G not assessed), indicating lower mortality (higher longevity). Shoot growth showed greater lime-induced stimulation (L > G,C) as compared to root growth. As a result the shoot:root ratio was higher in the limed rhizotrons than in the control (L > G,C). Liming induced a higher allocation of P, S, Mg, Ca and K to the leaves, stem and twigs. Gypsum showed similar effects, but was only significant for S. Liming increased the foliar Ca/Al ratio by both increasing foliar Ca and decreasing foliar Al, whereas gypsum did not clearly improve foliar nutrition. This study suggests that a moderate application of lime can be successful in stimulating seedling growth, but that gypsum had no effect on seedling growth. It can be concluded that this lime-induced growth stimulation is directly related to the improved soil fertility status, and the alleviation of Al toxicity and acid stress, resulting in better foliar nutrition. The impact of liming on fine roots, as a consequence, was not limited to a stimulation of the total amount of fine roots, but also improved the root uptake performance. This revised version was published online in June 2006 with corrections to the Cover Date.     

Biotechnical Characteristics of Root Systems of Typical Mediterranean Species

Quantifying patterns of fine root dynamics is crucial to the understanding of ecosystem structure and function, and in predicting how ecosystems respond to disturbance. Part of this understanding involves consideration of the carbon lost through root turnover. In the context of the rainfall pattern in the tropics, it was hypothesised that rainfall would strongly influence fine root biomass and longevity. A field study was conducted to determine root biomass, elemental composition and the influence of rainfall on longevity of fine roots in a tropical lowland evergreen rainforest at Danum Valley, Sabah, Malaysia. A combination of root coring, elemental analysis and rhizotron observation methods were used. Fine (less than 2 mm diameter) root biomass was relatively low (1700 kg ha −1) compared with previously described rainforest data. Standing root biomass was positively correlated with preceding rainfall, and the low fine root biomass in the dry season contained higher concentrations of N and lower concentrations of P and K than at other times. Observations on rhizotrons demonstrated that the decrease in fine root biomass in the dry season was a product of both a decrease in fine root length appearance and an increase in fine root length disappearance. Fitting an overall model to root survival time showed significant effects of rainfall preceding root disappearance, with the hazard of root disappearance decreasing by 8 for each 1 mm increase in the average daily (30 day) rainfall preceding root disappearance. While it is acknowledged that other factors have a part to play, this work demonstrates the importance of rainfall and soil moisture in influencing root biomass and root disappearance in this tropical rainforest.     

Using coloured roots to study root interaction and competition in intercropped legumes and non-legumes

Aims Root interactions between neighbour plants represent a fundamental aspect of the competitive dynamics in pure stand and mixed cropping systems. The comprehension of such phenomena places big methodological challenges, and still needs clarification. The objectives of this work were (i) to test if a species with coloured roots can be used to examine the interaction in a legume-non-legume intercropping system; (ii) to verify the importance of initial root growth on the successive root development of mixture component plants; (iii) to test if the root interaction in the shallow layers has consequences for deep root growth and (iv) to compare the effect of intraspecific and interspecific competition on root development and biomass growth.Methods A detailed study on root growth and interaction was carried out using rhizotron tubes where two legume species were grown in pure stands or were intercropped with red beet, a variety of Beta vulgaris L. with clear red roots. Within the rhizotrons, the three species were grown either without competitors, with two plants of the same species to measure intraspecific competition or with one legume and one red beet plant to study interspecific competition. The use of mixtures where one component has clearly coloured roots, together with several scalar measurements of root depth and proliferation, allowed the measurement of the root system of each species when grown in the mixtures.Important findings The use of rhizotron tubes coupled with species with coloured roots represented a valuable method to study the belowground interaction in mixed cropping systems. The initial root growth was a very important feature for the subsequent dominance of a species and it was not related to seed dimension. Initial root growth was also important because the root interactions in the shallower soil layers were found to influence the root growth in deeper soil. The root system of the red beet showed much faster and deeper growth than that of the legumes, and made red beet the dominant component in the mixtures while the legume root system was confined to the shallower soil layer. Intraspecific competition was well tolerated by the legumes, but it was limiting for the highly competitive red beet. The outcome of root interaction between neighbour plants was confirmed to be species-specific as it changed according to the intensity of the competitive effect/response of each species of the mixture: both legumes were slightly affected by the intraspecific and highly affected by interspecific competition while red beet was more affected by intraspecific competition but strongly dominant when intercropped with legumes.     

Nitrogen fixation of lucerne (Medicago sativa) in an acid soil: The use of rhizotrons as a model system to simulate field conditions

The effect of pelleting seeds of lucerne with lime was studied in an acid sandy soil. In pot experiments, the fraction of seedlings with crown nodules, i.e. nodules on the upper 10 mm of the taproot, increased from 26% to 71%. In rhizotrons, the application of CaCO₃ resulted in an even stronger response.An agar-contact method was used to study pH changes in the rhizosphere during a period up to 12 days. Application of 1.0 µmol of CaCO₃, in drops of 12 µL volume, resulted in an initial soil pH of 6.1 and yielded 75% crown nodulation. In the absence of CaCO₃, roots induced a pH increase from 5.1 (day 0) to 5.7 (day 12). However, this did not increase nodulation (5%). Obviously, this type of alkalinization does not overcome the acid-sensitive step of the nodulation process.