Summary 1In water-limited environments, the availability of water and nutrients to plants depends on environmental conditions, sizes and shapes of their root systems, and root competition. The goal of this study was to predict root system sizes and shapes for different plant growth forms using data on above-ground plant sizes, climate and soil texture. 2 A new data set of > 1300 records of root system sizes for individual plants was collected from the literature for deserts, scrublands, grasslands and savannas with ≤ 1000 mm mean annual precipitation (MAP). Maximum rooting depths, maximum lateral root spreads and their ratios were measured. 3 Root system sizes differed among growth forms and increased with above-ground size: annuals < perennial forbs = grasses < semi-shrubs < shrubs < trees. Stem succulents were as shallowly rooted as annuals but had lateral root spreads similar to shrubs. 4 Absolute rooting depths increased with MAP in all growth forms except shrubs and trees, but were not strongly related to potential evapotranspiration (PET). Except in trees, root systems tended to be shallower and wider in dry and hot climates and deeper and narrower in cold and wet climates. Shrubs were more shallowly rooted under climates with summer than winter precipitation regimes. 5 Relative to above-ground plant sizes, root system sizes decreased with increasing PET for all growth forms, but decreased with increasing MAP only for herbaceous plants. Thus relative rooting depths tended to increase with aridity, although absolute rooting depths decreased with aridity. 6 Using an independent data set of 20 test locations, rooting depths were predicted from MAP using regression models for three broad growth forms. The models succeeded in explaining 62% of the observed variance in median rooting depths. 7 Based on the data analysed here, Walter's two-layer model of soil depth partitioning between woody and herbaceous plants appears to be most appropriate in drier regimes (< 500 mm MAP) and in systems with substantial winter precipitation.
Studies in global plant biogeography have almost exclusively analyzed relationships of abiotic and biotic factors with the distribution and structure of vegetation aboveground. The goal of this study was to extend such analyses to the belowground structure of vegetation by determining the biotic and abiotic factors that influence vertical root distributions in the soil, including soil, climate, and plant properties. The analysis used a database of vertical root profiles from the literature with 475 profiles from 209 geographic locations. Since most profiles were not sampled to the maximum rooting depth, several techniques were used to estimate the amount of roots at greater depths, to a maximum of 3 m in some systems. The accuracy of extrapolations was tested using a subset of deeply (>2 m) sampled or completely sampled profiles. Vertical root distributions for each profile were characterized by the interpolated 50% and 95% rooting depths (the depths above which 50% or 95% of all roots were located). General linear models incorporating plant life‐form dominance, climate, and soil variables explained as much as 50% of the variance in rooting depths for various biomes and life‐forms. Annual potential evapotranspiration (PET) and precipitation together accounted for the largest proportion of the variance (12–16% globally and 38% in some systems). Mean 95% rooting depths increased with decreasing latitude from 80° to 30° but showed no clear trend in the tropics. Annual PET, annual precipitation, and length of the warm season were all positively correlated with rooting depths. Rooting depths in tropical vegetation were only weakly correlated with climatic variables but were strongly correlated with sampling depths, suggesting that even after extrapolation, sampling depths there were often insufficient to characterize root profiles. Globally, >90% of all profiles had at least 50% of all roots in the upper 0.3 m of the soil profile (including organic horizons) and 95% of all roots in the upper 2 m. Deeper rooting depths were mainly found in water‐limited ecosystems. Deeper 95% rooting depths were also found for shrublands compared to grasslands, in sandy soils vs. clay or loam soils, and in systems with relatively shallow organic horizons compared with deeper organic horizons.
Summary1 Root competition is defined as a reduction in the availability of a soil resource to roots that is caused by other roots. Resource availability to competitors can be affected through resource depletion (scramble competition) and by mechanisms that inhibit access of other roots to resources (contest competition, such as allelopathy). 2 It has been proposed that soil heterogeneity can cause size-asymmetric root competition. Support for this hypothesis is limited and contradictory, possibly because resource uptake is affected more by the amount and spatial distribution of resource-acquiring organs, relative to the spatial distribution of resources, than by root system size per se . 3 Root competition intensity between individual plants generally decreases as resource availability (but not necessarily habitat productivity) increases, but the importance of root competition relative to other factors that structure communities may increase with resource availability. 4 Soil organisms play important, and often species-specific, roles in root interactions. 5 The findings that some roots can detect other roots, or inert objects, before they are contacted and can distinguish between self and non-self roots create experimental challenges for those attempting to untangle the effects of self/non-self root recognition, self-inhibition and root segregation or proliferation in response to competition. Recent studies suggesting that root competition may represent a 'tragedy-of-the-commons' may have failed to account for this complexity. 6 Theories about potential effects of root competition on plant diversity (and vice versa) appear to be ahead of the experimental evidence, with only one study documenting different effects of root competition on plant diversity under different levels of resource availability. 7 Roots can interact with their biotic and abiotic environments using a large variety of often species-specific mechanisms, far beyond the traditional view that plants interact mainly through resource depletion. Research on root interactions between exotic invasives and native species holds great promise for a better understanding of the way in which root competition may affect community structure and plant diversity, and may create new insights into coevolution of plants, their competitors and the soil community.
The concept of a zone of influence, the area over which a plant alters the environment, forms the basis of many models of plant competition. Because of logistical difficulties, we actually know little about the sizes and shapes of zones of influence belowground. Here we advocate obtaining data on plants' belowground zones of influence, including the length and distribution of lateral roots, in order to understand better how plants respond to their abiotic soil environment and to other plants. We provide several examples from recent work. First, we present an analysis of a large global data set which shows that maximum lateral root spread correlates with canopy size but that, for a given canopy size, maximum lateral root spread is greater in arid environments and in coarse textured soils. Second, we use an experiment with the weedy annual Abutilon theophrasti to show how using nutrient analogs as tracers yields information about lateral root distributions within populations. In our experimental populations, the belowground zone of influence extended well beyond the closest neighboring plants. Overlap in zones of influence increased in nutrient patches. Third, we propose a new conceptual model of belowground zones of influence based on these and other data sets. The model assumes that the probability of resource uptake or competing with a particular neighbor declines with distance from the stem but that considerable uptake at great distances from the stem is still possible. It also allows for plasticity in root distributions as might occur in spatially heterogeneous soils. Finally, we suggest how better information on the shapes and sizes of belowground zones of influence will help develop a more predictive framework for understanding plant competition.
Pit membranes in bordered pits between neighbouring vessels play a major role in the entry of air-water menisci from an embolised vessel into a water-filled vessel (i.e., air-seeding). Here, we investigate intervessel pit membrane thickness (TPM) and embolism resistance (P50, i.e., the water potential corresponding to 50% loss of hydraulic conductivity) across a broad range of woody angiosperm species. Data on TPM and double intervessel wall thickness (TVW) were compiled based on electron and light microscopy. Fresh material that was directly fixated for transmission electron microscopy (TEM) was investigated for 71 species, while non-fresh samples were frozen, stored in alcohol, or air dried prior to TEM preparation for an additional 60 species. TPM and P50 were based on novel observations and literature. A strong correlation between TPM and P50 was found for measurements based on freshly fixated material (r = 0.78, P >0.01, n = 37), and between TPM and TVW (r = 0.79, P >0.01, n = 59), while a slightly weaker relationship occurred between TVW and P50 (r = 0.40, P >0.01, n = 34). However, non-fresh samples showed no correlation between TPM and P50, and between TPM and TVW. Intervessel pit membranes in non-fresh samples were c.28% thinner and more electron dense than fresh samples. Our findings demonstrate that TPM measured on freshly fixated material provides one of the strongest wood anatomical correlates of droughtinduced embolism resistance in angiosperms. Assuming that cellulose microfibrils show an equal spatial density, TPM is suggested to affect the length and the shape of intervessel pit membrane pores, but not the actual pore size. Moreover, the shrinking effect observed for TPM after dehydration and frost is associated with an increase in microfibril density and porosity, which may provide a functional explanation for embolism fatigue.
Vascular plants transport water under negative pressure without constantly creating gas bubbles that would disable their hydraulic systems. Attempts to replicate this feat in artificial systems almost invariably result in bubble formation, except under highly controlled conditions with pure water and only hydrophilic surfaces present. In theory, conditions in the xylem should favor bubble nucleation even more: there are millions of conduits with at least some hydrophobic surfaces, and xylem sap is saturated or sometimes supersaturated with atmospheric gas and may contain surface-active molecules that can lower surface tension. So how do plants transport water under negative pressure? Here, we show that angiosperm xylem contains abundant hydrophobic surfaces as well as insoluble lipid surfactants, including phospholipids, and proteins, a composition similar to pulmonary surfactants. Lipid surfactants were found in xylem sap and as nanoparticles under transmission electron microscopy in pores of intervessel pit membranes and deposited on vessel wall surfaces. Nanoparticles observed in xylem sap via nanoparticle-tracking analysis included surfactant-coated nanobubbles when examined by freeze-fracture electron microscopy. Based on their fracture behavior, this technique is able to distinguish between dense-core particles, liquid-filled, bilayer-coated vesicles/liposomes, and gas-filled bubbles. Xylem surfactants showed strong surface activity that reduces surface tension to low values when concentrated as they are in pit membrane pores. We hypothesize that xylem surfactants support water transport under negative pressure as explained by the cohesion-tension theory by coating hydrophobic surfaces and nanobubbles, thereby keeping the latter below the critical size at which bubbles would expand to form embolisms.
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