The drivers underlying the development of deep root systems, whether genetic or environmental, are poorly understood but evidence has accumulated that deep rooting could be a more widespread and important trait among plants than commonly anticipated from their share of root biomass. Even though a distinct classification of “deep roots” is missing to date, deep roots provide important functions for individual plants such as nutrient and water uptake but can also shape plant communities by hydraulic lift (HL). Subterranean fauna and microbial communities are highly influenced by resources provided in the deep rhizosphere and deep roots can influence soil pedogenesis and carbon storage.Despite recent technological advances, the study of deep roots and their rhizosphere remains inherently time-consuming, technically demanding and costly, which explains why deep roots have yet to be given the attention they deserve. While state-of-the-art technologies are promising for laboratory studies involving relatively small soil volumes, they remain of limited use for the in situ observation of deep roots. Thus, basic techniques such as destructive sampling or observations at transparent interfaces with the soil (e.g., root windows) which have been known and used for decades to observe roots near the soil surface, must be adapted to the specific requirements of deep root observation. In this review, we successively address major physical, biogeochemical and ecological functions of deep roots to emphasize the significance of deep roots and to illustrate the yet limited knowledge. In the second part we describe the main methodological options to observe and measure deep roots, providing researchers interested in the field of deep root/rhizosphere studies with a comprehensive overview. Addressed methodologies are: excavations, trenches and soil coring approaches, minirhizotrons (MR), access shafts, caves and mines, and indirect approaches such as tracer-based techniques.

1. There is a fundamental trade-off between leaf traits associated with either resource acquisition or resource conservation. This gradient of trait variation, called the economics spectrum, also applies to fine roots, but whether it is consistent for coarse roots or at the plant community level remains untested. 2. We measured a set of morphological and chemical root traits at a community level (functional parameters; FP) in 20 plant communities located along land-use intensity gradients and across three climatic zones (tropical, mediterranean and montane). We hypothesized (i) the existence of a root economics spectrum in plant communities consistent within root types (fine, < 2 mm; coarse, 2-5 mm), (ii) that variations in root FP occur with soil depths (top 20 cm of soil and 100-150 cm deep) and (iii) along land-use gradients. 3. Root FP covaried, in line with the resource acquisition-conservation trade-off, from communities with root FP associated with resource acquisition (e.g. high specific root length, SRL; thin diameters and low root dry matter contents, RDMC) to root FP associated with resource conservation (e.g. low SRL, thick diameters and high RDMC). This pattern was consistent for both fine and coarse roots indicating a strong consistency of a trade-off between resource acquisition and conservation for plant roots. 4. Roots had different suites of traits at different depths, suggesting a disparity in root function and exploitation capacities. Shallow, fine roots were thinner, richer in nitrogen and with lower lignin concentrations associated with greater exploitation capacities compared to deep, fine roots. Shallow, coarse roots were richer in nitrogen, carbon and soluble concentrations than deep, coarse roots. 5. Fine root parameters of highly disturbed, herbaceous-dominated plant communities in poorer soils were associated with foraging strategies, that is greater SRL and lower RDMC and lignin concentration than those from less disturbed communities. Coarse roots, however, were less sensitive to the land-use gradient. 6. Synthesis. This study demonstrates the existence of a general trade-off in root construction at a community level, which operates within all root types, suggesting that all plant tissues are controlled by the trade-off between resource acquisition and conservation.

Current knowledge gaps are identified and new lines of research for improving our understanding of the processes that drive deep root growth and functioning are proposed. This ultimately leads to a reflection on an alternative paradigm that could be used in the future as a unifying framework to describe and analyse deep rooting. Despite the many hurdles that pave the way to a practical understanding of deep rooting functions, it is anticipated that, in the relatively near future, increased knowledge about the deep rooting traits of a variety of plants and crops will have direct and tangible influence on how we manage natural and cultivated ecosystems.

No abstract

SummaryIrrigation by surge flooding does not always wet the soils thoroughly, and we have investigated the reasons for this on an irrigated plot in northern Senegal by monitoring the water budget during a rice cropping season (100 days). The amount of water added during each irrigation event was measured, and evapotranspiration and infiltration were estimated with lysimeters and Muntz infiltration rings, respectively. At the same time, piezometric levels, neutron probe values and water tension data were recorded at two stations in the plot. These measurements showed unusual results: infiltration rate was less than 1 Â 10 À6 mm s À1 (less than 0.1 mm a day), there was a constant water deficit during the entire irrigation period, around 50 cm deep, and tensiometers at 40 cm reacted very slowly to water infiltration. The water fluxes in the vadose zone derived from these data showed clearly a discrepancy between fluxes calculated from hydraulic gradients and fluxes calculated from mass conservation. The hydraulic gradients suggested a zero flux plane at 40 cm below the surface, but the calculated values of the fluxes overestimated by several orders of magnitude the infiltration rates determined on the plot, whereas fluxes determined from mass conservation matched far better. These results show that air was entrapped between the shallow water table and the wetting front, and this inhibited water infiltration. Modelling water flow down the soil profile with a computer program for simulating one-dimensional water movement (Hydrus) confirmed that single-phase models cannot describe imbibition in this situation. Simple infiltration models based on a modified Green-Ampt equation accounting for air compression and air counterflow, however, fit experimental infiltration data much better. We demonstrated that where surge flooding is associated with a shallow water table, as in many large irrigation schemes, one must take into account the presence of air to quantify the flow of water into the soil.

Melioidosis, a severe infection with the environmental bacterium Burkholderia pseudomallei, is being recognised increasingly frequently. What determines its uneven distribution within endemic areas is poorly understood. We cultured soil from a rice field in Laos for B. pseudomallei at different depths on 4 occasions over a 13-month period. We also measured physical and chemical parameters in order to identify associated characteristics. Overall, 195 of 653 samples (29.7%) yielded B. pseudomallei. A higher prevalence of B. pseudomallei was found at soil depths greater than the 30 cm currently recommended for B. pseudomallei environmental sampling. B. pseudomallei was associated with a high soil water content and low total nitrogen, carbon and organic matter content. Our results suggested that a sampling grid of 25 five metre square quadrats (i.e. 25 × 25 m) should be sufficient to detect B. pseudomallei at a given location if samples are taken at a soil depth of at least 60 cm. However, culture of B. pseudomallei in environmental samples is difficult and liable to variation. Future studies should both rely on molecular approaches and address the micro-heterogeneity of soil when investigating physico-chemical associations with the presence of B. pseudomallei.

No abstract

Fine root dynamics is a main driver of soil carbon stocks, particularly in tropical forests, yet major uncertainties still surround estimates of fine root production and turnover. This lack of knowledge is largely due to the fact that studying root dynamics in situ, particularly deep in the soil, remains highly challenging. We explored the interactions between fine root dynamics, soil depth, and rainfall in mature rubber trees (Hevea brasiliensis Müll. Arg.) exposed to sub-optimal edaphic and climatic conditions. A root observation access well was installed in northern Thailand to monitor root dynamics along a 4.5 m deep soil profile. Image-based measurements of root elongation and lifespan of individual roots were carried out at monthly intervals over 3 years. Soil depth was found to have a significant effect on root turnover. Surprisingly, root turnover increased with soil depth and root half-life was 16, 6–8, and only 4 months at 0.5, 1.0, 2.5, and 3.0 m deep, respectively (with the exception of roots at 4.5 m which had a half-life similar to that found between depths of 1.0 and 2.5 m). Within the first two meters of the soil profile, the highest rates of root emergence occurred about 3 months after the onset of the rainy season, while deeper in the soil, root emergence was not linked to the rainfall pattern. Root emergence was limited during leaf flushing (between March and May), particularly within the first two meters of the profile. Between soil depths of 0.5 and 2.0 m, root mortality appeared independent of variations in root emergence, but below 2.0 m, peaks in root emergence and death were synchronized. Shallow parts of the root system were more responsive to rainfall than their deeper counterparts. Increased root emergence in deep soil toward the onset of the dry season could correspond to a drought acclimation mechanism, with the relative importance of deep water capture increasing once rainfall ceased. The considerable soil depth regularly explored by fine roots, even though significantly less than in surface layers in terms of root length density and biomass, will impact strongly the evaluation of soil carbon stocks.