Fibrous root turnover and growth in sugar beet (Beta vulgaris var. saccharifera) as affected by nitrogen shortage

The root system of plants is subject to fast cycles of renewal and decay within the growing season. In water and/or nutrient stress conditions, this turnover may become strategic for plant survival and productivity, but knowledge about its mechanisms is still insufficient. In order to investigate the effects of nitrogen fertilization on growth and turnover of sugar beet roots, an experiment was carried out over two growing seasons in northern Italy with two levels of N supply (0, 100 kg ha^–1). Biomass production and partitioning were followed during growth, and fibrous root dynamics were inspected by means of computer-aided procedures applied to minirhizotron images.In conditions of N shortage, lower yields (storage roots) were associated with greater allocation of biomass to tap roots (final tap-root/shoot ratio = 5.6 vs. 4.1) and shallower distribution of fibrous root length density. The maximum depth of roots was not affected by N, but unfertilized plants reached it more slowly.The ratio of cumulative root dead length to produced length at the end of the growing period (TDL _max/TPL _max) was used as the most suitable approach for assessing overall root turnover. This ratio was greater in controls (0.73 vs. 0.50), which showed lower root longevity (–34% life-span on average), indicating that a greater proportion of root growth was renewed by unfertilized plants over the season.     

Most probable numbers of Azospirillum associated with the roots of inoculated pearl millet

Summary Inoculation of pearl millet (Pennisetum americanum (L.) Leeke) with Azospirillum significantly increased the numbers of this organism in the rhizosphere, rhizoplane, washed and crushed roots and surface sterilized and crushed roots. The maximum number of organisms plant^–1 were localized in the rhizosphere. The numbers of Azospirillum on the roots of inoculated plants grown under sterilized conditions were much higher than in the field grown plants. In both cases populations outside the roots were higher than in the surface sterilized roots. The highest numbers per unit root weight were recorded between 60–75 days of growth. N₂-ase activity throughout the growth cycle was very low and was not related to the populations of Azospirillum on the roots. Root exudates and extracts of pearl millet showed a stimulatory effect on the growth of Azospirillum suggesting their possible involvement in the colonization of this organism on the roots of inoculated plants.     

The effects of shading and N status on root proliferation in nutrient patches by the perennial grass Agropyron desertorum in the field

Competition for light can affect exploitation of spatially heterogeneous soil resources. To evaluate the influence of shoot status on root growth responses in nutrient-rich soil patches, we studied the effects of shading and whole-plant nitrogen status on root growth in N-enriched and nonenriched patches by mature Agropyron desertorum plants growing in the field with below-ground competition. Roots in enriched patches had greater length to weight ratios (specific root length, SRL), indicating increased absorptive surface areas, compared with roots in control patches. Increased SRL was due to increased production and length of higher order laterals rather than morphological changes in roots of the same branching order. Although the pattern of root growth rates in patches was the same for shaded and unshaded plants, the magnitude of this response to enriched patches was damped by shading. Root relative growth rates (RGR) in N-enriched patches were reduced by more than 50% by short-term shading treatments (60% reduction in photosynthetic flux density), while root RGR in unenriched patches was unaffected by shading. Unexpectedly, plants with higher nitrogen status had greater root RGR in enriched patches than plants that had not received nitrogen supplement, again with no detectable effect on root RGR in the unenriched patches. Therefore, while both shading and plant N status affected the ability of roots to exploit enriched patches by proliferation, there was no stimulation or suppression of root growth in the unenriched, control patches. Thus, plants already under competitive pressure above ground for light and below ground for nutrients should be less able to rapidly respond to opportunities presented in nutrient patches and pulses.     

Seasonal variation in the growth and nutrient content of yellow-birch replacement roots

Summary Modified air layers were established on lateral long roots of 9 yellow birch (Betula alleghniensis Britton) trees, and all replacement roots >. 5 cm long were harvested periodically during the 1971 and 1972 growing seasons. The first replacement roots grew 6 weeks after layer establishment. Root layers were inactive from 29 Oct. 71 to 5 May 72. Active root layers produced an average of 208 mg per tree during the first season and 198 mg per tree during the second season. Concentrations of N, P, K, Ca, Mg, Fe, Mn, Zn, and Al all varied within growing season, and average concentration of some elements—Ca in particular—varied between growing seasons. This technique shows promise for studying the nutrient status of root systems of forest trees.     

Impact of soil stockpiling on ericoid mycorrhizal colonization and growth of velvetleaf blueberry (Vaccinium myrtilloides) and Labrador tea (Ledum groenlandicum)

Soil stockpiling is a common practice prior to the reclamation of surface mines. In this study, velvetleaf blueberry and Labrador tea plants were grown from seed in fresh soil, stockpiled soil (1 year), and autoclaved stockpiled soil (1 year) obtained from the Canadian boreal forest. After 7 months of growth, the root colonization intensity with ericoid mycorrhizal (ERM) fungi in both plants growing in stockpiled soil was lower compared to plants growing in the fresh soil. The diversity of ERM fungal species in roots also decreased due to soil stockpiling and Pezoloma ericae was absent from the plants growing in stockpiled soil. Changes in the ERM root colonization in plants growing in stockpiled soil were accompanied by decreases in root and shoot dry weights. Leaf chlorophyll, nitrogen, and phosphorus concentrations of velvetleaf blueberry were higher in fresh soil compared to 1‐year stockpiled soil. Plants grown in the autoclaved stockpiled soil became colonized by the thermotolerant ERM fungus Leohumicola verrucosa and showed higher root and shoot biomass compared to the nonautoclaved stockpiled soil. The results point to the importance of ERM fungi for growth of ericaceous plants, even under favorable environmental conditions and adequate fertilization, and suggest that reduced ERM colonization intensity and ERM fungal diversity in roots likely contributed to the negative effects of soil stockpiling on growth of velvetleaf blueberry and Labrador tea.     

An in situ approach for measuring root-associated respiration and nitrate uptake of forest trees

The ecophysiological characteristics of fine roots of mature forest plants are poorly understood because of difficulties of measurement. We explored a root in-growth approach to measure respiration and nitrate uptake of woody plant roots in situ. Roots of seven species were grown into sand-filled chambers. Root-associated respiration was measured as CO ₂ emission on four dates and nitrate uptake was quantified using ¹⁵N. All the roots were younger than 3 months at the time of measurement. Fine root respiration measured over the temperature range of 14.5–15.5 °C averaged 18.9–36.5 nmol gDM ^–1 s ^–1 across species. Nitrate uptake rates by these fine roots (1.3–6.8 nmol gDM ^–1 s ^–1) were comparable to other studies of forest trees. The root respiration rates were several times higher than measurements on detached roots of mature trees, concurring with literature observations that young roots respire much more rapidly than older roots. The root in-growth approach appears promising for providing information on the metabolic activity of fine roots of mature forest trees growing in soil.     

Impact of galactoglucomannan oligosaccharides on elongation growth in intact mung bean plants

The impact of exogenously applied galactoglucomannan oligosaccharides (GGMOs) and their structurally modified forms (GGMOs-r—galactoglucomannosyl alditols, GGMOs-g—with reduced galactose content) on the growth of mung bean (Vigna radiata (L.) Wilczek) intact plants cultured in hydroponics has been determined. GGMOs alone or in combination with exogenously added IBA have influenced (with stimulation and/or inhibition effect) hypocotyl and seminal root elongation, adventitious and lateral roots formation and elongation in dependency on their concentration used. The inhibition of elongation growth in hypocotyls as well as in roots was connected with changes of cell wall-associated peroxidases activity and is probably associated with the beginning of cell wall rigidification. Data presented in this paper confirm the hypothesis that exogenously added GGMOs may have antiauxin activity and may interact also with endogenous growth regulators. Certain monosaccharide sequences with terminal galactose in the side chain of GGMOs probably play important role in their biological activity in intact plants as it was demonstrated previously in individual parts of plants.     

The production of a phytotoxin,nectrolide, by some root-surface isolates ofCylindrocarpon radicicola,Wr.

Summary A phytotoxic compound, for which the name nectrolide has been proposed, was isolated fromCylindrocarpon radicicola. Analysis of nectrolide and its physical and chemical properties show that the compound is identical with brefeldin A, an antibiotic produced byPenicillium brefeldianum.Twenty different isolates ofC. radicicola from plant roots were examined for their ability to produce nectrolide in liquid culture. Twelve of these yielded nectrolide at concentrations varying from 31 to 82 µg per ml when grown for 14 days on Czapek-Dox liquid medium. Eight isolates did not yield the metabolite in culture.At a concentration of 6 µg per ml nectrolide arrested the growth of germinated blackbutt seedlings, whilst at lower concentrations (2–4 µg per ml) the compound caused severe stunting and blackening of the roots of seedling plants. Nectrolide watered onto blackbutt seedlings growing in forest soils can be detected in the roots after 24 hrs. The compound has been crystallised from ether extracts of tomato roots supplied with pure nectrolide and can be detected in stem tissues within 24 hours of watering it onto the roots.As a result of the widespread association ofC. radicicola with unthrifty plants and the phytotoxic nature of nectrolide, it is suggested that this compound may function in a number of plant diseases, the causes of which have not been elucidated.     

Timing of root dormancy and tolerance to root waterlogging in clonal Sitka spruce

 Actively growing root tips of Picea sitchensis (Bong.) Carr. plants are highly susceptible to damage if waterlogged, but they are known to have some tolerance after they stop growing in the autumn. This paper describes the selection of clones on the basis of root dormancy timing and the corresponding responses of their roots to over-winter waterlogging. Sitka spruce transplants of Alaska, Queen Charlotte Islands (QCI), and Washington provenances were screened for early or late root dormancy over 2 successive years. Cuttings were propagated from the selected plants and after growing on for 2 years, they were planted in transparent acrylic tubes within outdoor ‘root observation chambers’. Extension of main roots and the timing of onset of root dormancy was recorded on the clonal plants. The tubes were flooded in November and maintained with a water table 280 mm below the soil surface until March of the next year. Waterlogging caused most main root tips to die back, but within 2 months of draining regeneration occurred on the main roots below the waterlogging level. This regeneration was most commonly the growth of existing lateral tips or production of new lateral roots. Roots of early-dormant Washington plants died back on average 129 mm less than late-dormant Washington plants, and early-dormant Alaska plants had 173 mm less dieback than late-dormant Alaska plants. Differences between the clones of the QCI provenance were not significant. The 40% and 52% increases in survival depth of roots in early-dormant Washington and Alaska clones respectively indicates a potential for improving the rooting depth of Sitka spruce on seasonally waterlogged soils by planting clones selected on the basis of root dormancy. Received: 14 July 1997 / Accepted: 15 September 1997     

Factors affecting formation of cluster roots in Myrica gale seedlings in water culture

The effect of nutrient deficiency, aeration, phosphorus supply, and nitrogen source on the formation of cluster (proteoid) roots was examined in Myrica gale seedlings growing in water culture. Only the omission of phosphorus resulted in the formation of significant numbers to cluster roots when plants were grown in a number of 1/4 strength Hoagland's solutions, each lacking one mineral nutrient. Aeration shortened the time required for cluster root formation and increased the percentage of plants forming cluster roots. The proportion of the root system comprised of cluster roots decreased as the phosphorus concentration in the solution increased and no cluster roots formed in solutions containing 8 mg P/L. Phosphorus supply also affected total plant biomass, proportion of biomass comprising nitrogen-fixing nodules, shoot:root ratio, phosphorus concentration in the leaves and phosphorus content of the plants. The plants showed luxury consumption of phosphorus and were able to produce large amounts of biomass utilizing only stored phosphorus.Nitrogen source also affected cluster root formation. Urea-fed plants produced cluster roots more quickly and devoted a substantially larger proportion of root growth to cluster roots than did nitrate-fed plants. The longest cluster root axes were produced in nitrate-fed plants supplied with no phosphorus and the shortest were in urea-fed plants at 4 mg P L^–1.Four methods for expressing the extent of cluster root formation were examined and it was concluded that cluster roots as a proportion of total fine root dry weight is preferable in many cases. Formation of cluster roots in response to phosphorus deficiency coupled with previously demonstrated traits allows Myrica gale to adapt to a wide range of soil conditions.     

The morphological changes of wheat genotypes as affected by the levels of localized phosphate supply

Effects of localized phosphate supply on the seedling growth of wheat (Triticum aestivum L.) genotypes 81(85)5-3-3-3 (P-efficient) and NC37 (P-inefficient) were studied using a device which allowed only 3 cm length of root segment to be exposed to phosphate treatment. Localized supply of 0 mmol P L^–1 and the rest of root supplied with 0.1 mmol P L^–1 (HLH), increased the shoot height, leaf area, root/shoot ratio for 81(85)5-3-3-3, length of root and root axis for NC37, and root axis length and density of first-laterals for both the genotypes, compared to plants with the whole root system in P-sufficient solution (HHH). This suggested that above- and below-ground morphological parameters of wheat were promoted by a localized P-deficiency, presumably via a P deficiency signal. There was a significant difference in the number of first-order laterals between the two wheat genotypes when most of the roots were grown without P and only 3 cm length of root was supplied with 0.3 mmol P L^–1. The relationship between the number and density of 2nd-order lateral roots and level of local P supply was quadratic. Maximum number and density of 2nd-order lateral roots were obtained with a localized P supply of 0.70 mmol L^–1.     

Colonization potential of bacteria in the rhizosphere

The effect of inoculum density on growth and steady-state populations of aPseudomonas sp., aMycoplana sp., and aCurtobacterium sp. in the rhizosphere if gnotobiotic barley plants was studied. Inoculation of sterile barley seedling at concentrations of about 1×10³, 1×10⁵ and 1×10⁷ viable cells (mg dry wt root)^–1 resulted in rapid colonization; maximum populations of about 5×10⁷ viable cells (mg dry wt root)^–1 developed in each case. We define this maximum population as the colonization potential. Measurement of growth of known rhizosphere bacteria might be a useful index of the amount of available carbon and energy lost by growing roots.     

Role of kinetin in alleviation of copper and zinc toxicity in Lupinus termis plants

The effect of exogenous kinetin application on the growth and some physiological processes of Lupinus termis plants growing in metal containing solutions with excess concentrations of Cu and Zn ion were studied. Generally, plants growing in these solutions had a lower chlorophyll (Chl.) content, leaf relative water content (RWC) and produced less biomass than the control plants. Proline content was higher in metal-treated plants than in untreated controls. Chromatography of cell-free-extracts of roots and shoots indicated three main protein peaks with molecular weights about 170, 75--70 and 5--45 kDa. These peaks were coincident with Cu or Zn maxima. Addition of kinetin reduced the decline in Chl. content in metal-treated plants, improved water status of the plants and enhanced growth of the shoots and roots. The Cu or Zn content expressed on a per mg protein basis was raised when kinetin was applied to the growing shoots. Kinetin (Kin), Cu and Zn, singly and in the presence of kinetin (Cu × Kin and Zn × Kin), significantly affected the parameters tested. Only the effects of Cu × Kin and Zn × Kin interactions on shoot fresh weight and Cu × Kin on root length were statistically insignificant. Based on the calculated coefficient of determination ( ²) the roles of Cu and Zn in affecting Chl. content and growth were dominant in comparison to kinetin. Kinetin effect was dominant for root length and proline content, but the role of the interaction was subdominant. The results of this study indicate that kinetin can alleviate the harmful effects of Cu and Zn on the growth of lupin plants through stimulation of Cu and Zn incorporation into metal-binding proteins.     

Morphology, ecology, and contaminant removal efficiency of eight wetland plants with differing root systems

The objective of this study was to test the hypothesis that fibrous-root plants and rhizomatic-root plants are characterized by different root morphologies, root growth and distribution, and contaminant removal capabilities. Four fibrous-root and four rhizomatic-root wetland plants were studied in mono-cultured microcosms which received wastewater. Fibrous-root plants had significantly greater (P < 0.05) small-size root (diameter ≤ 1 mm) biomass and a larger (P < 0.05) root surface area per plant than the rhizomatic-root plants and exhibited accelerated growth in both shoots and roots compared to the rhizomatic-root plants. Fibrous-root plants developed the majority of their root biomass increment within a shallower gravel medium than the rhizomatic-root plants. All plants demonstrated fast root biomass growth from July to September. The wetland microcosms planted with fibrous-root plants showed significantly higher (P < 0.05) ammonium-nitrogen (NH₄-N) and nitrate-nitrogen (NO₃-N) removal rates from July to December than those planted with the rhizomatic-root plants. These results suggest that root characteristics of wetland plants, which are related to their shoot and root growth, root distribution, and decontamination ability, can be used in the selection of wetland plants with a higher contaminant removal capacity and in the construction of a multi-species wetland plant community. Handling editor: S. M. Thomaz     

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.     

Hyperaccumulation of Lead by Roots, Hypocotyls, and Shoots of Brassica juncea

The effects of different concentrations of lead nitrate (10^–5 to 10^–3 M) on root, hypocotyl, and shoot growth of Indian mustard (Brassica juncea L. var. megarrhiza), and the uptake and accumulation of Pb²⁺ by its roots, hypocotyls, and shoots were investigated. Lead had no significant inhibitory effect on the root growth at concentrations of 10^–5 to 10^–4 M during the entire treatment, while at 10^–3 M, Pb slightly inhibited the root and shoot growth. B. juncea has ability to take up Pb from solutions and accumulate it in its roots, and transport and concentrate it. The Pb contents in the parts of plants treated with 10^–3 M Pb were greater than those of untreated plants, by factors of 230 in the roots, 170 in the hypocotyls, and 3 in the shoots.     

Root exudates as mediators of mineral acquisition in low-nutrient environments

Plant developmental processes are controlled by internal signals that depend on the adequate supply of mineral nutrients by soil to roots. Thus, the availability of nutrient elements can be a major constraint to plant growth in many environments of the world, especially the tropics where soils are extremely low in nutrients. Plants take up most mineral nutrients through the rhizosphere where micro-organisms interact with plant products in root exudates. Plant root exudates consist of a complex mixture of organic acid anions, phytosiderophores, sugars, vitamins, amino acids, purines, nucleosides, inorganic ions (e.g. HCO₃ ^–, OH^–, H⁺), gaseous molecules (CO₂, H₂), enzymes and root border cells which have major direct or indirect effects on the acquisition of mineral nutrients required for plant growth. Phenolics and aldonic acids exuded directly by roots of N₂-fixing legumes serve as major signals to Rhizobiaceae bacteria which form root nodules where N₂ is reduced to ammonia. Some of the same compounds affect development of mycorrhizal fungi that are crucial for phosphate uptake. Plants growing in low-nutrient environments also employ root exudates in ways other than as symbiotic signals to soil microbes involved in nutrient procurement. Extracellular enzymes release P from organic compounds, and several types of molecules increase iron availability through chelation. Organic acids from root exudates can solubilize unavailable soil Ca, Fe and Al phosphates. Plants growing on nitrate generally maintain electronic neutrality by releasing an excess of anions, including hydroxyl ions. Legumes, which can grow well without nitrate through the benefits of N₂ reduction in the root nodules, must release a net excess of protons. These protons can markedly lower rhizosphere pH and decrease the availability of some mineral nutrients as well as the effective functioning of some soil bacteria, such as the rhizobial bacteria themselves. Thus, environments which are naturally very acidic can pose a challenge to nutrient acquisition by plant roots, and threaten the survival of many beneficial microbes including the roots themselves. A few plants such as Rooibos tea (Aspalathus linearis L.) actively modify their rhizosphere pH by extruding OH^– and HCO₃ ^– to facilitate growth in low pH soils (pH 3 – 5). Our current understanding of how plants use root exudates to modify rhizosphere pH and the potential benefits associated with such processes are assessed in this review.     

Growth responses to waterlogging and drainage of woody Hakea (Proteaceae) seedlings,originating from contrasting habitats in south-western Australia

We investigated the responses of seven woody Hakea (Proteaceae) species (two populations of each), to two months of waterlogging and subsequent drainage, in a controlled glasshouse experiment. The species originated from contrasting environments (winter-wetland versus non-wetland habitats), and differed in abundance (endangered ironstone species versus common species). Waterlogging arrested growth of the main root system, and stimulated the formation of superficial adventitious roots just below the root/shoot junction in all species. Wetland species produced at least twice the amount of adventitious root dry mass of that of non-wetland species, due to differences in number, length or degree of branching. Their adventitious roots also tended to have higher porosities (7–10% versus 5–6% gas spaces). The relative amount of adventitious roots formed was strongly, positively correlated with the maintenance of shoot growth, and only the non-wetland species showed significant shoot growth reductions (19%) upon waterlogging. Dry mass percentage of stems and leaves, and leaf dry mass per area (LMA) increased considerably during waterlogging in all species (averages of 15, 29 and 27%, respectively), returning to the values of continuously drained control plants after drainage. Similarly, upon drainage, a suppression of shoot growth (average 35%) and a stimulation of root growth (average 50%) restored the root mass ratios to those of control plants. We found a negative correlation between the maintenance of growth during waterlogging versus that after waterlogging, suggesting a trade-off in functioning of the superficial adventitious roots between waterlogged and drained conditions. The rare winter-wet ironstone endemics resembled the common winter-wet species in most of their responses to waterlogging and drainage. Therefore, the results presented here cannot offer an explanation for their different distribution patterns. Our results suggest that non-wetland species may be disadvantaged in a wetland environment, due to their lower capacity to form adventitious roots resulting in stronger growth reductions.     

Root growth dynamics of Deyeuxia angustifolia seedlings in response to water level

A greenhouse experiment was performed to investigate root growth dynamics, plant growth, root porosity and root morphology of a marsh plant Deyeuxia angustifolia, one of the dominant species in the Sanjiang Plain, China. The aim of this study was to elucidate how this plant adjusts its root system to acclimate to different hydrological environments. Experimental treatments included three water depths: −5, 0 and 5 cm (relative to the soil surface). Biomass accumulation was higher in the −5 cm (0.90 g per plant) and 0 cm water-depth (1.18 g per plant) than that in the 5 cm water-depth treatments (0.66 g per plant), indicating that plant growth was inhibited in the high water level. Root:shoot ratio (0.67 versus 0.42–0.43), the length (16 cm versus 12–13 cm) and diameter of adventitious roots (0.47 mm versus 0.41 mm), and root number (167 versus 81–119 number of roots per plant) were higher in the 0 cm water-depth than those in the high and low water-depth treatments. Enhanced water level led to slightly increased porosity of main roots, but porosity was about 7% in all treatments. After 8 weeks, roots had been distributed into 14, 11 and 7 cm soil depth in the 0, −5 and 5 cm water-depth treatments, respectively, indicating that both high and low water levels led to shallow root systems. Our data suggest that D. angustifolia can adjust root morphology and root growth pattern according to water level, and that this plant has limited oxygen diffusion potential to the roots due to the reduced biomass in the high water level.     

Mycorrhizal C costs and nutritional benefits in developing grapevines

Arbuscular mycorrhizal (AM) C-costs in grapevines were investigated. Dormant vines rely on stored C for initial growth. Therefore AM colonisation costs would compete with plant growth for available C reserves. One-year-old grapevines, colonised with Glomus etunicatum (Becker and Gerdemann), were cultivated under glasshouse conditions. The C-economy and P utilisation of the symbiosis were sequentially analysed. AM colonisation, during the 0–67 day growth period, used more stem C relative to root C, which resulted in lower shoot growth. The decline in AM colonisation during the period of 67–119 days coincided with stem C replenishment and higher shoot growth. Construction costs of AM plants and root C allocation increased with root P uptake. The efficiency of P utilisation was lower in AM roots. The reliance of AM colonisation on stem C declined with a decrease in colonisation, providing more C for the refilling of stem carbohydrate reserves and shoot growth. Once established, the AM symbiosis increased P uptake at the expense of refilling of root C reserves. Although higher root C allocation increased plant construction costs, AM roots were more efficient at P utilisation.