Improving rice tolerance to potassium deficiency by enhancing OsHAK16p:WOX11‐controlled root development

Potassium (K) deficiency in plants confines root growth and decreases root‐to‐shoot ratio, thus limiting root K acquisition in culture medium. A WUSCHEL‐related homeobox (WOX) gene, WOX11, has been reported as an integrator of auxin and cytokinin signalling that regulates root cell proliferation. Here, we report that ectopic expression of WOX11 gene driven by the promoter of OsHAK16 encoding a low‐K‐enhanced K transporter led to an extensive root system and adventitious roots and more effective tiller numbers in rice. The WOX11‐regulated root and shoot phenotypes in the OsHAK16p:WOX11 transgenic lines were supported by K‐deficiency‐enhanced expression of several RR genes encoding type‐A cytokinin‐responsive regulators, PIN genes encoding auxin transporters and Aux/IAA genes. In comparison with WT, the transgenic lines showed increases in root biomass, root activity and K concentrations in the whole plants, and higher soluble sugar concentrations in roots particularly under low K supply condition. The improvement of sugar partitioning to the roots by the expression of OsHAK16p:WOX11 was further indicated by increasing the expression of OsSUT1 and OsSUT4 genes in leaf blades and several OsMSTs genes in roots. Expression of OsHAK16p:WOX11 in the rice grown in moderate K‐deficient soil increased total K uptake by 72% and grain yield by 24%–32%. The results suggest that enlarging root growth and development by the expression of WOX11 in roots could provide a useful option for increasing K acquisition efficiency and cereal crop productivity in low K soil.     

Low host specificity of root‐associated fungi at an Arctic site

In High Arctic ecosystems, plant growth and reproduction are limited by low soil moisture and nutrient availability, low soil and air temperatures, and a short growing season. Mycorrhizal associations facilitate plant nutrient acquisition and water uptake and may therefore be particularly ecologically important in nutrition‐poor and dry environments, such as parts of the Arctic. Similarly, endophytic root associates are thought to play a protective role, increasing plants' stress tolerance, and likely have an important ecosystem function. Despite the importance of these root‐associated fungi, little is known about their host specificity in the Arctic. We investigated the host specificity of root‐associated fungi in the common, widely distributed arctic plant species Bistorta vivipara, Salix polaris and Dryas octopetala in the High Arctic archipelago Svalbard. High‐throughput sequencing of the internal transcribed spacer 1 (ITS1) amplified from whole root systems generated no evidence of host specificity and no spatial autocorrelation within two 3 m × 3 m sample plots. The lack of spatial structure at small spatial scales indicates that Common Mycelial Networks (CMNs) are rare in marginal arctic environments. Moreover, no significant differences in fungal OTU richness were observed across the three plant species, although their root system characteristics (size, biomass) differed considerably. Reasons for lack of host specificity could be that association with generalist fungi may allow arctic plants to more rapidly and easily colonize newly available habitats, and it may be favourable to establish symbiotic relationships with fungi possessing different physiological attributes.     

An in situ approach to detect tree root ecology: linking ground‐penetrating radar imaging to isotope‐derived water acquisition zones

Tree root distribution and activity are determinants of belowground competition. However, studying root response to environmental and management conditions remains logistically challenging. Methodologically, nondestructive in situ tree root ecology analysis has lagged. In this study, we tested a nondestructive approach to determine tree coarse root architecture and function of a perennial tree crop, Theobroma cacao L., at two edaphically contrasting sites (sandstone and phyllite–granite derived soils) in Ghana, West Africa. We detected coarse root vertical distribution using ground‐penetrating radar and root activity via soil water acquisition using isotopic matching of δ¹⁸O plant and soil signatures. Coarse roots were detected to a depth of 50 cm, however, intraspecifc coarse root vertical distribution was modified by edaphic conditions. Soil δ¹⁸O isotopic signature declined with depth, providing conditions for plant–soil δ¹⁸O isotopic matching. This pattern held only under sandstone conditions where water acquisition zones were identifiably narrow in the 10–20 cm depth but broader under phyllite–granite conditions, presumably due to resource patchiness. Detected coarse root count by depth and measured fine root density were strongly correlated as were detected coarse root count and identified water acquisition zones, thus validating root detection capability of ground‐penetrating radar, but exclusively on sandstone soils. This approach was able to characterize trends between intraspecific root architecture and edaphic‐dependent resource availability, however, limited by site conditions. This study successfully demonstrates a new approach for in situ root studies that moves beyond invasive point sampling to nondestructive detection of root architecture and function. We discuss the transfer of such an approach to answer root ecology questions in various tree‐based landscapes.     

Mesorhizobium ciceri LMS-1 expressing an exogenous 1-aminocyclopropane-1-carboxylate (ACC) deaminase increases its nodulation abilities and chickpea plant resistance to soil constraints

Aims: Our goal was to understand the symbiotic behaviour of a Mesorhizobium strain expressing an exogenous 1‐aminocyclopropane‐1‐carboxylate (ACC) deaminase, which was used as an inoculant of chickpea (Cicer arietinum) plants growing in soil. Methods and Results: Mesorhizobium ciceri LMS‐1 (pRKACC) was tested for its plant growth promotion abilities on two chickpea cultivars (ELMO and CHK3226) growing in nonsterilized soil that displayed biotic and abiotic constraints to plant growth. When compared to its wild‐type form, the M. ciceri LMS‐1 (pRKACC) strain showed an increased nodulation performance of c. 125 and 180% and increased nodule weight of c. 45 and 147% in chickpea cultivars ELMO and CHK3226, respectively. Mesorhizobium ciceri LMS‐1 (pRKACC) was also able to augment the total biomass of both chickpea plant cultivars by c. 45% and to reduce chickpea root rot disease susceptibility. Conclusions: The results obtained indicate that the production of ACC deaminase under free living conditions by Mesorhizobium strains increases the nodulation, plant growth abilities and biocontrol potential of these strains. Significance and Impact of the Study: This is the first study regarding the use of a transformed rhizobial strain expressing an exogenous ACC deaminase in different plant cultivars growing in soil. Hence, obtaining Mesorhizobium strains with high ACC deaminase activity is a matter of extreme importance for the development of inoculants for field applications.     

Root Development and Nutrient Uptake

Root system formation proceeds in close coordination with shoot growth. Accordingly, root growth and its functions are regulated tightly by the shoot through materials cycling between roots and shoots. A plant root system consists of different kinds of roots that differ in morphology and functions. The spatial configuration and distribution of these roots determine root system architecture in the soil, which in turn primarily regulates the acquisition of soil resources like nutrients and water. Morphological and physiological properties of each root and the concomitant tissues further affect nutrient uptake and transport, while the root traits that are related to such acquisition also depend on the kinds of nutrients and their mobility in the soil. In addition, mechanisms involved in the uptake and transport of mineral nutrients recently have been elucidated at the molecular level. A number of genes for acquisition and transport of various mineral nutrients have been identified in model plant systems such as Arabidopsis thaliana, and rice, and in other plant species. An integration of studies on nutrient behavior in soils and the morphological and physiological functions of root systems will further elucidate the mechanism of plant nutrient uptake and transport by roots, and offer a real possibility of genetically improving crop productivity in problem soils.

     

Approaches for enhancement of N2 fixation efficiency of chickpea (Cicer arietinum L.) under limiting nitrogen conditions

Chickpea (Cicer arietinum) is an important pulse crop in many countries in the world. The symbioses between chickpea and Mesorhizobia, which fix N₂ inside the root nodules, are of particular importance for chickpea's productivity. With the aim of enhancing symbiotic efficiency in chickpea, we compared the symbiotic efficiency of C‐15, Ch‐191 and CP‐36 strains of Mesorhizobium ciceri in association with the local elite chickpea cultivar ‘Bivanij’ as well as studied the mechanism underlying the improvement of N₂ fixation efficiency. Our data revealed that C‐15 strain manifested the most efficient N₂ fixation in comparison with Ch‐191 or CP‐36. This finding was supported by higher plant productivity and expression levels of the nifHDK genes in C‐15 nodules. Nodule specific activity was significantly higher in C‐15 combination, partially as a result of higher electron allocation to N₂ versus H⁺. Interestingly, a striking difference in nodule carbon and nitrogen composition was observed. Sucrose cleavage enzymes displayed comparatively lower activity in nodules established by either Ch‐191 or CP‐36. Organic acid formation, particularly that of malate, was remarkably higher in nodules induced by C‐15 strain. As a result, the best symbiotic efficiency observed with C‐15‐induced nodules was reflected in a higher concentration of the total and several major amino metabolites, namely asparagine, glutamine, glutamate and aspartate. Collectively, our findings demonstrated that the improved efficiency in chickpea symbiotic system, established with C‐15, was associated with the enhanced capacity of organic acid formation and the activities of the key enzymes connected to the nodule carbon and nitrogen metabolism.     

Shifts in symbiotic associations in plants capable of forming multiple root symbioses across a long‐term soil chronosequence

Changes in soil nutrient availability during long‐term ecosystem development influence the relative abundances of plant species with different nutrient‐acquisition strategies. These changes in strategies are observed at the community level, but whether they also occur within individual species remains unknown. Plant species forming multiple root symbioses with arbuscular mycorrhizal (AM) fungi, ectomycorrhizal (ECM) fungi, and nitrogen‐(N) fixing microorganisms provide valuable model systems to examine edaphic controls on symbioses related to nutrient acquisition, while simultaneously controlling for plant host identity. We grew two co‐occurring species, Acacia rostellifera (N₂‐fixing and dual AM and ECM symbioses) and Melaleuca systena (AM and ECM dual symbioses), in three soils of contrasting ages (c. 0.1, 1, and 120 ka) collected along a long‐term dune chronosequence in southwestern Australia. The soils differ in the type and strength of nutrient limitation, with primary productivity being limited by N (0.1 ka), co‐limited by N and phosphorus (P) (1 ka), and by P (120 ka). We hypothesized that (i) within‐species root colonization shifts from AM to ECM with increasing soil age, and that (ii) nodulation declines with increasing soil age, reflecting the shift from N to P limitation along the chronosequence. In both species, we observed a shift from AM to ECM root colonization with increasing soil age. In addition, nodulation in A. rostellifera declined with increasing soil age, consistent with a shift from N to P limitation. Shifts from AM to ECM root colonization reflect strengthening P limitation and an increasing proportion of total soil P in organic forms in older soils. This might occur because ECM fungi can access organic P via extracellular phosphatases, while AM fungi do not use organic P. Our results show that plants can shift their resource allocation to different root symbionts depending on nutrient availability during ecosystem development.     

The importance of root gravitropism for inter-root competition and phosphorus acquisition efficiency: results from a geometric simulation model

We have observed that low soil phosphorus availability alters the gravitropic response of basal roots in common bean (Phaseolus vulgaris L.), resulting in a shallower root system. In this study we use a geometric model to test the hypotheses that a shallower root system is a positive adaptive response to low soil P availability by (1) concentrating root foraging in surface soil horizons, which generally have the highest P availability, and (2) reducing spatial competition for P among roots of the same plant. The growth of nine root systems contrasting in gravitropic response over 320 h was simulated in SimRoot, a dynamic three-dimensional geometric model of root growth and architecture. Phosphorus acquisition and inter-root competition were estimated with Depzone, a program that dynamically models nutrient diffusion to roots. Shallower root systems had greater P acquisition per unit carbon cost than deeper root systems, especially in older root systems. This was due to greater inter-root competition in deeper root systems, as measured by the volume of overlapping P depletion zones. Inter-root competition for P was a significant fraction of total soil P depletion, and increased with increasing values of the P diffusion coefficient (De), with root age, and with increasing root gravitropism. In heterogenous soil having greater P availability in surface horizons, shallower root systems had greater P acquisition than deeper root systems, because of less inter-root competition as well as increased root foraging in the topsoil. Root P acquisition predicted by SimRoot was validated against values for bean P uptake in the field, with an r ² between observed and predicted values of 0.75. Our results support the hypothesis that altered gravitropic sensitivity in P-stressed roots, resulting in a shallower root system, is a positive adaptive response to low P availability by reducing inter-root competition within the same plant and by concentrating root activity in soil domains with the greatest P availability. This revised version was published online in August 2006 with corrections to the Cover Date.     

Oryza sativa Lysine‐Histidine‐type Transporter 1 functions in root uptake and root‐to‐shoot allocation of amino acids in rice

In agricultural soils, amino acids can represent vital nitrogen (N) sources for crop growth and yield. However, the molecular mechanisms underlying amino acid uptake and allocation are poorly understood in crop plants. This study shows that rice (Oryza sativa L.) roots can acquire aspartate at soil concentration, and that japonica subspecies take up this acidic amino acid 1.5‐fold more efficiently than indica subspecies. Genetic association analyses with 68 representative japonica or indica germplasms identified rice Lysine‐Histidine‐type Transporter 1 (OsLHT1) as a candidate gene associated with the aspartate uptake trait. When expressed in yeast, OsLHT1 supported cell growth on a broad spectrum of amino acids, and effectively transported aspartate, asparagine and glutamate. OsLHT1 is localized throughout the rice root, including root hairs, epidermis, cortex and stele, and to the leaf vasculature. Knockout of OsLHT1 in japonica resulted in reduced root uptake of amino acids. Furthermore, in ¹⁵N‐amino acid‐fed mutants versus wild‐type, a higher percentage of ¹⁵N remained in roots instead of being allocated to the shoot. ¹⁵N‐ammonium uptake and subsequently the delivery of root‐synthesized amino acids to Oslht1 shoots were also significantly decreased, which was accompanied by reduced shoot growth. These results together provide evidence that OsLHT1 functions in both root uptake and root to shoot allocation of a broad spectrum of amino acids in rice.     

The combined effects of a long‐term experimental drought and an extreme drought on the use of plant‐water sources in a Mediterranean forest

Vegetation in water‐limited ecosystems relies strongly on access to deep water reserves to withstand dry periods. Most of these ecosystems have shallow soils over deep groundwater reserves. Understanding the functioning and functional plasticity of species‐specific root systems and the patterns of or differences in the use of water sources under more frequent or intense droughts is therefore necessary to properly predict the responses of seasonally dry ecosystems to future climate. We used stable isotopes to investigate the seasonal patterns of water uptake by a sclerophyll forest on sloped terrain with shallow soils. We assessed the effect of a long‐term experimental drought (12 years) and the added impact of an extreme natural drought that produced widespread tree mortality and crown defoliation. The dominant species, Quercus ilex, Arbutus unedo and Phillyrea latifolia, all have dimorphic root systems enabling them to access different water sources in space and time. The plants extracted water mainly from the soil in the cold and wet seasons but increased their use of groundwater during the summer drought. Interestingly, the plants subjected to the long‐term experimental drought shifted water uptake toward deeper (10–35 cm) soil layers during the wet season and reduced groundwater uptake in summer, indicating plasticity in the functional distribution of fine roots that dampened the effect of our experimental drought over the long term. An extreme drought in 2011, however, further reduced the contribution of deep soil layers and groundwater to transpiration, which resulted in greater crown defoliation in the drought‐affected plants. This study suggests that extreme droughts aggravate moderate but persistent drier conditions (simulated by our manipulation) and may lead to the depletion of water from groundwater reservoirs and weathered bedrock, threatening the preservation of these Mediterranean ecosystems in their current structures and compositions.     

Effects of transgenic soybean on growth and phosphorus acquisition in mixed culture system

Transgenic soybean plants overexpressing the Arabidopsis purple acid phosphatase gene AtPAP15 (OXp) or the soybean expansin gene GmEXPB2 (OXe) can improve phosphorous (P) efficiency in pure culture by increasing Apase secretion or changing root morphology. In this study, soybean‐soybean mixed cultures were employed to illuminate P acquisition among plants in mixed stands of transgenic and wild‐type soybean. Our results showed that transgenic soybean plants were much more competitive, and had greater growth and P uptake than wild‐type soybean in mixed culture in both low P calcareous and acid soils. Furthermore, OXe plants had an advantage in calcareous soils when mixed with OXp, whereas the latter performed much better in acid soils. In soybean‐maize mixed culture, transgenic soybean had no impact on maize growth compared to controls in both acid and calcareous soils with different P conditions. As for soybean in mixed culture, OXp plants had no significant advantages regardless of P availability or soil type, while P efficiency improved in OXe in calcareous soils compared to controls. These results imply that physiological traits could be easily affected by the mixed maize. Transgenic soybean plants with enhanced root traits had more competitive advantages than those with improved root physiology in mixed culture.     

脲酶抑制剂/硝化抑制剂对植稻土壤中尿素N行为的影响

采用自制根盒试验,主要研究了脲酶抑制剂氢醌（HQ）,硝化抑制剂双氰胺（DCD）及二者组合对离水稻根际不同距离处NH4^--N和NO3^-N分布的影响,结果表明,DCD及其与HQ组合均能显著促进稻株地上部分生长,始终显著降低水稻根际与近根际土中NH4^ -N含量直至施肥后60d,施肥后20d时,DCD及其与HQ组合可使非根际土中NH4^ -N含量显著增加,随后,却出现相反现象,施肥后20d时,距根际不同距离的土壤中,配施DCD或DCD＋HQ处理均能显著降低NO3^-N含量,随后,近根际和非根际仍保持上述现象直至施肥后40d;同未施DCD处理相比,根际土壤却较早出现NO3^--N含量高峰,正好与水稻N营养需求时期相一致,因此,DCD及其与HQ组合可减少水稻根际环境下尿素N损失潜势,通过不种稻土壤和距根际3cm处的土壤中尿素无机氮形态分布的差异,充分显示了研究水稻根际土壤氮素转化及相关抑制剂对其影响时,以取离根际3cm外的土壤作为非根际明显优于不种稻土壤。     

Sensitivity of chickpea and faba bean to root‐zone hypoxia,elevated ethylene,and carbon dioxide

During soil waterlogging, plants experience O₂ deficits, elevated ethylene, and high CO₂ in the root‐zone. The effects on chickpea (Cicer arietinum L.) and faba bean (Vicia faba L.) of ethylene (2 μL L^?1), CO₂ (2–20% v/v) or deoxygenated stagnant solution were evaluated. Ethylene and high CO₂ reduced root growth of both species, but O₂ deficiency had the most damaging effect and especially so for chickpea. Chickpea suffered root tip death when in deoxygenated stagnant solution. High CO₂ inhibited root respiration and reduced growth, whereas sugars accumulated in root tips, of both species. Gas‐filled porosity of the basal portion of the primary root of faba bean (23%, v/v) was greater than for chickpea (10%), and internal O₂ movement was more prominent in faba bean when in an O₂‐free medium. Ethylene treatment increased the porosity of roots. The damaging effects of low O₂, such as death of root tips, resulted in poor recovery of root growth upon reaeration. In conclusion, ethylene and high CO₂ partially inhibited root extension in both species, but low O₂ in deoxygenated stagnant solution had the most damaging effect, even causing death of root tips in chickpea, which was more sensitive to the low O₂ condition than faba bean.     

Effects of Resource-Island Soils, Competition, and Inoculation with Azospirillum on Survival and Growth of Pachycereus pringlei, the Giant Cactus of the Sonoran Desert

Resource‐island soils formed by some plants in arid lands are capable of supporting certain plants that do not normally establish in surrounding areas free of vegetation. We determined growth responses of Pachycereus pringlei (cardon), the giant columnar cactus, whose widespread, finely branched, subsurface root systems stabilize desert soils, to four soils collected from within or outside of resource islands. Traits of cardon grown in soils from mature (MM) or young (YM) Prosopis articulata (mesquite), mature Olnea testosa (ironwood, MI), or bare areas (BA) were compared, to determine differences between the effects of soils due to the identity or the stage of development of nurse plants. The levels of soil N, P, and C contents were in the order MM> > YM > MI BA. The BA soil had the coarsest and MM soil the finest texture. Cardon was also grown in pot cultures inoculated with the plant‐growth‐promoting bacterium Azospirillum brasilense, or in association with a competing grass, Sorghum bicolor (sorghum). Competition did not affect survival rates of cardon in any of the soils after six months of growth, but decreased biomass accumulation by up to 90% in the best (MM) soil. Inoculation of cardon seeds with A. brasilense did not affect survival but resulted in significantly better root and shoot growth, and this effect increased linearly as soil nutrients declined. In the best soil (MM), A. brasilense had no effect on cardon growth, but in the poorest soil (BA) shoot dry mass was almost 60% and root length over 100% greater as a result of inoculation, with responses in the other two soils intermediate. This effect did not appear to be owing to N₂ fixation, as nitrogenase activity (acetylene reduction) was not detected in any of the treatments. Soil formation by selected nurse trees in arid areas is an important factor in plant establishment and growth, and the present results indicate that these processes can be impeded or facilitated by the introduction of competing or beneficial organisms. The use of beneficial microorganisms associated with roots may accelerate the restoration of disturbed areas. Key words: Azospirillum, desert soils, nitrogen fixation, nurse plant, Pachycereus pringlei, plant survival, resource‐island.     

Assessment of the root system of Brassica juncea (L.) czern. and Bidens pilosa L. exposed to lead polluted soils using rhizobox systems

The purpose of this study was to compare the behavior of the root system of one of the most frequently cited species in phytoremediation Indian mustard [Brassica juncea (L.) Czern.] and a representative perennial herb (Bidens pilosa L.) native of Argentina, for different concentrations of lead in soils through chemical and visualization techniques of the rhizosphere. Lead polluted soils from the vicinity of a lead recycling plant in the locality of Bouwer, were used in juxtaposed rhizobox systems planted with seedlings of B. juncea and B. pilosa with homogeneous and heterogeneous soil treatments. Root development, pH changes in the rhizosphere, dry weight biomass, lead content of root and aerial parts and potential extraction of lead by rhizosphere exudates were determined. In both species lead was mainly accumulated in roots. However, although B. juncea accumulated more lead than B. pilosa at elevated concentrations in soils, the latter achieved greater root and aerial development. No changes in the pH of the rhizosphere associated to lead were observed, despite different extractive potentials of lead in the exudates of the species analyzed. Our results indicated that Indian mustard did not behave as a hyperaccumulator in the conditions of the present study.     

Comparative analysis of microbial diversity and bacterial seedling disease‐suppressive activity in organic‐farmed and standardized commercial conventional soils for rice nursery cultivation

The outbreak of rice plant diseases can be effectively suppressed in organic farming systems. However, the mechanisms of disease suppression by organic farming systems are not well understood. When Burkholderia‐infected rice seeds were sown and cultivated on nine organic‐farmed soils which were supplied by nine independent organic rice farmers or standardized commercial conventional soils, the emergence of bacterial seedling diseases was suppressed to equivalent degrees in nine organic‐farmed soils, whereas the diseases occurred in two commercial conventional soils. In any organic or commercial conventional soil sown with healthy rice seeds as a control, the diseases did not appear. Upon physicochemical analysis of the nine organic‐farmed soils, component common to these organic‐farmed soils seemed to not be directly associated with disease‐suppressive activity. However, microbiome analyses indicated that the bacterial population in these nine organic‐farmed soils was more diverse than those in commercial conventional soils. Intriguingly, the diverse bacterial population structures of organic‐farmed soils were preserved after irrigating and sowing rice seeds, but that of commercial conventional soils was clearly changed by them. Thus, organic‐farmed soils seem to maintain robust bacterial populations despite the irrigation and seedling growth. Indeed, pathogenic Burkholderia in infected rice seeds also did not proliferate in the seedling grown on organic‐farmed soils. Taken together, the common feature of organic‐farmed soils might be the correlation between bacterial seedling disease‐suppressive activity and higher robustness of the diversified microbiome.     

Plant litter chemistry alters the content and composition of organic carbon associated with soil mineral and aggregate fractions in invaded ecosystems

Through the input of disproportionate quantities of chemically distinct litter, invasive plants may potentially influence the fate of organic matter associated with soil mineral and aggregate fractions in some of the ecosystems they invade. Although context dependent, these native ecosystems subjected to prolonged invasion by exotic plants may be instrumental in distinguishing the role of plant–microbe–mineral interactions from the broader edaphic and climatic influences on the formation of soil organic matter (SOM). We hypothesized that the soils subjected to prolonged invasion by an exotic plant that input recalcitrant litter (Japanese knotweed, Polygonum cuspidatum) would have a greater proportion of plant‐derived carbon (C) in the aggregate fractions, as compared with that in adjacent soil inhabited by native vegetation that input labile litter, whereas the soils under an invader that input labile litter (kudzu, Pueraria lobata) would have a greater proportion of microbial‐derived C in the silt‐clay fraction, as compared with that in adjacent soils that receive recalcitrant litter. At the knotweed site, the higher C content in soils under P. cuspidatum, compared with noninvaded soils inhabited by grasses and forbs, was limited to the macroaggregate fraction, which was abundant in plant biomarkers. The noninvaded soils at this site had a higher abundance of lignins in mineral and microaggregate fractions and suberin in the macroaggregate fraction, partly because of the greater root density of the native species, which might have had an overriding influence on the chemistry of the above‐ground litter input. At the kudzu site, soils under P. lobata had lower C content across all size fractions at a 0–5 cm soil depth despite receiving similar amounts of Pinus litter. Contrary to our prediction, the noninvaded soils receiving recalcitrant Pinus litter had a similar abundance of plant biomarkers across both mineral and aggregate fractions, potentially because of the higher surface area of soil minerals at this site. The plant biomarkers were lower in the aggregate fractions of the P. lobata‐invaded soils, compared with noninvaded pine stands, potentially suggesting a microbial co‐metabolism of pine‐derived compounds. These results highlight the complex interactions among litter chemistry, soil biota, and minerals in mediating soil C storage in unmanaged ecosystems; these interactions are particularly important under global changes that may alter plant species composition and hence the quantity and chemistry of litter inputs in terrestrial ecosystems.     

Architectural Tradeoffs between Adventitious and Basal Roots for Phosphorus Acquisition

Adventitious rooting contributes to efficient phosphorus acquisition by enhancing topsoil foraging. However, metabolic investment in adventitious roots may retard the development of other root classes such as basal roots, which are also important for phosphorus acquisition. In this study we quantitatively assessed the potential effects of adventitious rooting on basal root growth and whole plant phosphorus acquisition in young bean plants. The geometric simulation model SimRoot was used to dynamically model root systems with varying architecture and C availability growing for 21 days at 3 planting depths in 3 soil types with contrasting nutrient mobility. Simulated root architectures, tradeoffs between adventitious and basal root growth, and phosphorus acquisition were validated with empirical measurements. Phosphorus acquisition and phosphorus acquisition efficiency (defined as mol phosphorus acquired per mol C allocated to roots) were estimated for plants growing in soil in which phosphorus availability was uniform with depth or was greatest in the topsoil, as occurs in most natural soils. Phosphorus acquisition and acquisition efficiency increased with increasing allocation to adventitious roots in stratified soil, due to increased phosphorus depletion of surface soil. In uniform soil, increased adventitious rooting decreased phosphorus acquisition by reducing the growth of lateral roots arising from the tap root and basal roots. The benefit of adventitious roots for phosphorus acquisition was dependent on the specific respiration rate of adventitious roots as well as on whether overall C allocation to root growth was increased, as occurs in plants under phosphorus stress, or was lower, as observed in unstressed plants. In stratified soil, adventitious rooting reduced the growth of tap and basal lateral roots, yet phosphorus acquisition increased by up to 10% when total C allocation to roots was high and adventitious root respiration was similar to that in basal roots. With C allocation to roots decreased by 38%, adventitious roots still increased phosphorus acquisition by 5%. Allocation to adventitious roots enhanced phosphorus acquisition and efficiency as long as the specific respiration of adventitious roots was similar to that of basal roots and less than twice that of tap roots. When adventitious roots were assigned greater specific respiration rates, increased adventitious rooting reduced phosphorus acquisition and efficiency by diverting carbohydrate from other root types. Varying the phosphorus diffusion coefficient to reflect varying mobilities in different soil types had little effect on the value of adventitious rooting for phosphorus acquisition. Adventitious roots benefited plants regardless of basal root growth angle. Seed planting depth only affected phosphorus uptake and efficiency when seed was planted below the high phosphorus surface stratum. Our results confirm the importance of root respiration in nutrient foraging strategies, and demonstrate functional tradeoffs among distinct components of the root system. These results will be useful in developing ideotypes for more nutrient efficient crops.