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Generalizations of the Moran effect explaining spatial synchrony in population fluctuations.

The Moran effect for populations separated in space states that the autocorrelations in the population fluctuations equal the autocorrelation in environmental noise, assuming the same linear density regulation in all populations. Here we generalize the Moran effect to include also nonlinear density regulation with spatial heterogeneity in local population dynamics as well as in the effects of environmental covariates by deriving a simple expression for the correlation between the sizes of two populations, using diffusion approximation to the theta-logistic model. In general, spatial variation in parameters describing the dynamics reduces population synchrony. We also show that the contribution of a covariate to spatial synchrony depends strongly on spatial heterogeneity in the covariate or in its effect on local dynamics. These analyses show exactly how spatial environmental covariation can synchronize fluctuations of spatially segregated populations with no interchange of individuals even if the dynamics are nonlinear.

Demography↗

Temporal and spatial variation of episodic wind erosion in unburned and burned semiarid shrubland.

Redistribution of soil, nutrients, and contaminants is often driven by wind erosion in semiarid shrublands. Wind erosion depends on wind velocity (particularly during episodic, high-velocity winds) and on vegetation, which is generally sparse and spatially heterogeneous in semiarid ecosystems. Further, the vegetation cover can be rapidly and greatly altered due to disturbances, particularly fire. Few studies, however, have evaluated key temporal and spatial components of wind erosion with respect to (i) erosion rates on the scale of weeks as a function of episodic high-velocity winds, (ii) rates at unburned and burned sites, and (iii) within-site spatial heterogeneity in erosion. Measuring wind erosion in unburned and recently burned Chihuahuan desert shrubland, we found (i) weekly wind erosion was related more to daily peak wind velocities than to daily average velocities as consistent with our findings of a threshold wind velocity at approximately 7 m s(-1); (ii) greater erodibility in burned vs. unburned shrubland as indicated by erosion thresholds, aerodynamic roughness, and nearground soil movement; and (iii) burned shrubland lost soil from intercanopy and especially canopy patches in contrast to unburned shrubland, where soil accumulated in canopy patches. Our results are among the first to quantify post-fire wind erosion and highlight the importance of accounting for finer temporal and spatial variation in shrubland wind erosion. This finer-scale variation relates to semiarid land degradation, and is particularly relevant for predictions of contaminant resuspension and redistribution, both of which historically ignore finer-scale temporal and spatial variation in wind erosion.

Conservation of Natural Resources↗

Oscillations of two competing microbial populations in configurations of two interconnected chemostats.

It is known that, when two microbial populations competing for a single rate-limiting nutrient are grown in a spatially uniform environment, such as a single chemostat, with competition being the only interaction between them, they cannot coexist, but eventually one of the two populations prevails and the other becomes extinct. Spatial heterogeneity has been suggested as a means of obtaining coexistence of the two populations. A configuration of two interconnected chemostats is a simple model of a spatially heterogeneous environment. It has been shown that, when Monod's model is used for the specific growth rates of the two populations, steady-state coexistence can be obtained in such systems for wide ranges of operating conditions. In the present work, we study a model of microbial competition in configurations of interconnected chemostats and we show that, if a substrate inhibition model is used for the specific growth rates of the two populations, coexistence in a periodic state is also possible. The analysis of the model is done by numerical bifurcation theory methods.

Environmental Microbiology↗

Tracking cardiac electrical instability by computing interlead heterogeneity of T-wave morphology.

Oscillations in T-wave morphology, particularly T-wave alternans (TWA), have been fundamentally linked to increased susceptibility to ventricular fibrillation (VF). We investigated whether the escalation in complexity of T-wave oscillations before VF is attributable to increased spatial heterogeneity of repolarization. Peak interlead T-wave heterogeneity (TWH) was measured by second central moment analysis of T-wave morphology in epicardial electrograms in dogs during left anterior descending coronary artery occlusion. TWH differentiated cases in which myocardial ischemia provoked VF from those without VF (563 +/- 56 vs. 139 +/- 36 microV, P < 0.01). In the former group, progressive, significant increases in TWH above preocclusion baseline (70 +/- 8 microV) began at 2.25 min after the start of occlusion and were associated successively with TWA (at 155 +/- 19 microV), T-wave multupling (at 386 +/- 100 microV), complex oscillatory T-wave forms (at 560 +/- 76 microV), discordant TWA (at 572 +/- 98 microV), and VF at 4.36 +/- 0.14 min. TWH in precordial ECGs in 12 pigs during angioplasty-balloon-induced myocardial ischemia also discriminated animals that experienced VF (from 90 +/- 14 at baseline to 382 +/- 39 microV, P < 0.05) from those without VF (from 96 +/- 17 at baseline to 199 +/- 61 microV, NS). Ischemia-induced changes in ST segment and T-wave amplitude did not predict VF. Heightened spatial heterogeneity of repolarization, as assessed by second central moment analysis of TWH, underlies TWA and increased risk for ischemia-induced VF. Monitoring spatial TWH from precordial leads could prove useful in stratifying risk for life-threatening arrhythmias.

Algorithms↗

Effects of pulmonary blood flow on the fractal nature of flow heterogeneity in sheep lungs.

The spatial heterogeneity of pulmonary blood flow can be described by the relative dispersion (RD) of weight-flow histograms (RD = SD/mean). Glenny and Robertson (J. Appl. Physiol. 69: 532-545, 1990) showed that RD of flow in the lung is fractal in nature, characterized by the fractal dimension (D) and RD for the smallest realizable volume element (RDref). We studied the effects of increasing total pulmonary blood flow on D and RDref. In eight in situ perfused sheep lung preparations, 15-microns radio-labeled microspheres were injected into the pulmonary artery at five different blood flows ranging, in random order, from 1.5 to 5.0 l/m. The lungs were in zone 2 at the lower flows and in zone 3 at the higher flows. The lungs were removed, dried, cut into 2 x 2 x 2-cm3 pieces, weighed, and then counted for microsphere radioactivity. Fractal plots of log(weight) vs. log(RD) were constructed by iteratively combining neighboring pieces and then calculating RD with the increasingly larger portion size. D, which is one minus the slope of the fit through this plot, was 1.14 +/- 0.09 and did not change as blood flow increased. However, RDref decreased significantly (P < 0.01) as total flow increased. We conclude that the fractal nature of pulmonary blood flow distribution is not altered by changes in overall flow.

Animals↗

Heterogeneity of microvascular response to ischemia in skeletal muscle.

The aim of this study was to use the video-microscopic approach to analyze simultaneously, at very low magnification, red cell velocity (VRBC) and density of perfused capillaries (CD) in an entire capillary network of a frog sartorius muscle before and after 3 s, 1 min and 30 min arterial occlusions. In 11 muscles studied, the 3 s occlusion caused a significant increase in the overall peak hyperemic VRBC and CD (from 0.14 +/- 0.08 to 0.28 +/- 0.16 S.D. mm/s, and from 104 +/- 41 to 126 +/- 42 S.D. cap/mm2). All muscles participated in this overall response. On average, 71% of capillaries per network showed hyperemia. The 1 min occlusion also caused significant increases in overall VRBC and CD (from 0.14 +/- 0.08 to 0.34 +/- 0.17 S.D. mm/s, and from 94 +/- 44 to 123 +/- 44 S.D. cal/mm2). In contrast to the 3 s response, only 9 out of 11 muscles participated in this response, with an average capillary participation of 80%. The 30 min occlusion caused a significant increase in overall VRBC (from 0.13 +/- 0.07 to 0.24 +/- 0.17 S.D. mm/s) but no change in CD. Only 6 out of 11 muscles participated in this response, with an average capillary participation of 56%. In terms of these microvascular data, the present study demonstrate for the first time ever (1) an appreciable spatial heterogeneity of responses within the capillary network in a single muscle following all three types of occlusions, and (2) a progressively larger muscle-to-muscle heterogeneity of responses with longer duration of occlusion. Since the spatial heterogeneity (i.e. increase, no change or even decrease in flow) was associated with groups of similarly behaving capillaries supplied by a common arteriole, the existence of a flow-suppressing arteriolar mechanism (in addition to metabolic and myogenic vasodilating mechanisms) is postulated to explain the dramatic heterogeneity of response to ischemia in this muscle.

Animals↗

[Spatial pattern of lithorheophile macrobenthos (by the example of a stream in Moscow region)].

The distribution of lithorheophil macrobenthos was studied in the range of scales from decimeter (regions in the scope of one stone) up to seven kilometers on the basis of 153 quantitative samples from oligosaprobic stream in Moscow region. Community shows significant spatial heterogeneity even at the small scale: life forms inhabit different sides of each stone and different sites of the riffle depending on current. At the scale of tens and hundreds of meters (between different riffles, within one stream) spatial heterogeneity of benthos is low, distribution of common species is independent from each other and riffs are similar by community structure. At the scope of kilometers heterogeneity increase and the alterations of community appear due to changes in stream size. Seasonal differences of benthos distribution are mentioned. The impact of various environmental factors on the distribution of lithorheophile macrobenthos is discussed.

Animals↗

Spatial variability in the mineralisation of the phenylurea herbicide linuron within a Danish agricultural field: multivariate correlation to simple soil parameters.

The spatial variability in the mineralisation rate of linuron [N-(3,4-dichlorophenyl)-N'-methoxy-N'-methylurea] was studied within a previously treated Danish agricultural field by sampling soils from eleven different plots randomly distributed across an area of 20 x 20 m. The soils were characterised with respect to different abiotic and biotic properties including moisture content, organic matter content, pH, nutrient content, bacterial biomass, potential for mineralisation of MCPA [(4-chloro-2-methylphenoxy)acetic acid] and linuron. Five soils had a potential for mineralisation of linuron, with 5-15% of the added [ring-U-14C]linuron metabolised to 14CO2 within 60 days at 10 degrees C, while no extensive mineralisation of linuron was observed in the six remaining soils within this period. A TLC analysis of the methanol-extractable residues showed no development of 14C-labelled metabolites from linuron in any of the samples. Multivariate analysis was conducted to elucidate relationships between the intrinsic properties of single soil samples and initial rate of linuron mineralisation. The analysis indicated that important soil parameters in determining the spatial heterogeneity included the C(total)/N(total) ratio, pH and the water-extractable potassium contents, with the first of these highly negatively correlated and the last two highly positively correlated to the initial linuron mineralisation rate. This study shows that enhanced biodegradation of linuron may develop with successive field treatments, but that considerable in-field spatial heterogeneity in the degradation rate still exists. Combined with a parallel enrichment study focused on the underlying microbial processes, the present results suggest that intrinsic soil properties affect the linuron-metabolising bacterial population and thereby determine the spatial variability in the linuron mineralisation activity.

Biodegradation, Environmental↗

Radiomics as a spatial context for treatment decision-making in head and neck cancer.

Radiomics has been widely explored as a non-invasive biomarker in head and neck squamous cell carcinoma (HNSCC), yet its clinical role remains unclear. Tissue-based biomarkers differ in their susceptibility to spatial sampling. Biomarkers such as PD-L1 expression, immune-cell infiltration, necrosis, and immune exclusion may exhibit substantial spatial heterogeneity, whereas HPV/p16 status and some genomic alterations are generally more stable across the tumor. Nevertheless, localized sampling may incompletely capture heterogeneity in selected clinical contexts. This mismatch becomes clinically relevant when treatment decisions, particularly for chemoradiotherapy, immunotherapy, or de-escalation, are based on potentially non-representative biopsy findings. In this narrative review, we argue that the role of radiomics is not to outperform established biomarkers, but to contextualize them by capturing spatial heterogeneity related to hypoxia, necrosis, stromal architecture, and immune exclusion. We synthesize current evidence linking radiomic features to these biological processes and map them to specific clinical decision points, including larynx preservation, immunotherapy stratification, and recurrence assessment. Rather than serving as a standalone predictor, radiomics may provide complementary spatial information that helps identify situations in which biopsy-derived biomarkers should be interpreted with caution. Although current evidence is largely retrospective, radiomics offers a pragmatic framework for integrating spatial information into biomarker-guided clinical workflows.

Journal Article↗

Oxi4503, a novel vascular targeting agent: effects on blood flow and antitumor activity in comparison to combretastatin A-4 phosphate.

Oxi4503, which is the diphosphate prodrug of combretastatin A1, is a novel vascular targeting agent from the combretastatin family. Another member of this family, Combretastatin A-4 phosphate (CA4P), is a well-characterized vascular targeting agent already being evaluated in clinical trials. The potential for tumor vascular targeting by Oxi4503 was assessed in a mouse system. This approach aims to shut down the established tumor vasculature, leading to the development of extensive tumor cell necrosis. The vascular effects of Oxi4503 were assessed in the s.c. implanted MDA-MB-231 adenocarcinoma and the MHEC5-T hemangio-endothelioma in SCID mice and in a range of normal tissues. Blood flow was measured by i.v. injection of fluorescence beads, while quantitative fluorescence microscopy was used to measure the spatial heterogeneity of blood flow in tumor sections. Oxi4503 induced the shutdown of tumor blood vessels in a dose-dependent pattern with an ED50 at 3 mg/kg in contrast to 43 mg/kg of CA4P. Quantitative fluorescence microscopy showed that Oxi4503 increased the spatial heterogeneity in tumor blood flow. Oxi4503 affected peripheral tumor regions less than central regions, although this was not as pronounced as seen with CA4P, where only central regions were affected. The vascular shutdown induced by administration of Oxi4503 at a dose of 6 mg/kg resulted in extensive cell loss 24 hours following treatment, which translated into a significant effect on tumor growth. Tumor growth was completely repressed at doses above 12.5 mg/kg of Oxi4503, while doses above 25 mg/kg showed tumor regression and even complete regression in some animals. These results are promising for the use of Oxi4503 as a tumor vascular targeting agent. Moreover the potent antitumor effect when administered as a single agent suggests a different activity profile than CA4P.

Adenocarcinoma↗

Preferential localization of active mitochondria in process tips of immature retinal oligodendrocytes.

Active mitochondria were visualized with a laser scanning confocal microscope using the dye rhodamine 123, which is accumulated in mitochondria with a high respiratory activity. In glial precursor cells cultured from rabbit retina the fluorescence signal of active mitochondria was confined to the tips of processes, while in oligodendrocytes an even distribution was found. The developmental stage of cultured oligodendrocytes was determined by staining with O4 and O10 antibodies after the end of microfluorometric measurements. In glial precursor cells, the mitochondrial uncoupler CCCP generated an increase in the cytoplasmic calcium concentration ([Ca2+]i) in regions corresponding to those revealed by rhodamine 123 staining. In mature oligodendrocytes CCCP evoked [Ca2+]i elevation without significant spatial heterogeneity. However, electron microscopy did not reveal an uneven distribution of mitochondria in either immature or mature oligodendrocytes. We conclude that in the precursor cells, active mitochondria are confined to process tips and serve as a Ca2+ pool, while in oligodendrocytes such a spatial heterogeneity cannot be detected. This preferential localization of active mitochondria may play a role in the neurone-glia interaction during the formation of the contact between the myelinating glial process and the axonal membrane.

Aniline Compounds↗

Patch dynamics and metapopulation theory: the case of successional species.

We present a mathematical framework that combines extinction-colonization dynamics with the dynamics of patch succession. We draw an analogy between the epidemiological categorization of individuals (infected, susceptible, latent and resistant) and the patch structure of a spatially heterogeneous landscape (occupied-suitable, empty-suitable, occupied-unsuitable and empty-unsuitable). This approach allows one to consider life-history attributes that influence persistence in patchy environments (e.g., longevity, colonization ability) in concert with extrinsic processes (e.g., disturbances, succession) that lead to spatial heterogeneity in patch suitability. It also allows the incorporation of seed banks and other dormant life forms, thus broadening patch occupancy dynamics to include sink habitats. We use the model to investigate how equilibrium patch occupancy is influenced by four critical parameters: colonization rate, extinction rate, disturbance frequency and the rate of habitat succession. This analysis leads to general predictions about how the temporal scaling of patch succession and extinction-colonization dynamics influences long-term persistence. We apply the model to herbaceous, early-successional species that inhabit open patches created by periodic disturbances. We predict the minimum disturbance frequency required for viable management of such species in the Florida scrub ecosystem.

Disasters↗

Double dissociation between the effects of peri-postrhinal cortex and hippocampal lesions on tests of object recognition and spatial memory: heterogeneity of function within the temporal lobe.

It is widely believed that declarative memory is mediated by a medial temporal lobe memory system consisting of several distinct structures, including the hippocampus and perirhinal cortex. The strong version of this view assumes a high degree of functional homogeneity and serial organization within the medial temporal lobe, such that double dissociations between individual structures should not be possible. In the present study, we tested for a functional double dissociation between the hippocampus and peri-postrhinal cortex in a single experiment. Rats with bilateral excitotoxic lesions of either the hippocampus or peri-postrhinal cortex were assessed in tests of spatial memory (radial maze) and object recognition memory. For the latter, the spontaneous object recognition task was conducted in a modified apparatus designed to minimize the potentially confounding influence of spatial and contextual factors. A clear functional double dissociation was observed: rats with hippocampal lesions were impaired relative to controls and those with peripostrhinal cortex lesions on the spatial memory task, whereas rats with peri-postrhinal lesions were impaired relative to the hippocampal and control groups in object recognition. These results provide strong evidence in favor of heterogeneity and independence of function within the temporal lobe.

Animals↗

Investigating the heterogeneity of cell growth in microbial colonies by FTIR microspectroscopy.

Microorganisms rarely occur as individual cells in nature and are, instead, organized in complex multicellular communities such as colonies, fruiting bodies, or biofilms. Interest in the natural microbial life-style has increased during the last decade and a whole plethora of techniques has been used to gain insight into the development, structure and composition of diverse microbial communities. We have developed a technique for investigating the spatial heterogeneity of microbial growth in macro-colonies which essentially entails excision of the colonies with the underlying agar, freezing and subsequent cryotoming of the colonies, then FTIR microspectroscopic mapping of the cryosections. Colonies from Legionella, Bacillus, and Candida strains were chosen as model systems of multi-cellular communities to evaluate the technique. The results obtained indicate pronounced cell population heterogeneity even in relatively young colonies cultivated under laboratory conditions. Spectral data obtained from different positions within, e.g., a colony of Legionella bozemanii 120 h old indicated that levels of the storage material poly-beta-hydroxybutyric acid were significantly higher in cells at the surface of the colonies than in those growing at the bottom next to the agar surface. Similarly, in a 24-h-old macro-colony of Bacillus megaterium significantly more of the capsular compound polyglutamic acid was detected in upper layers than in deeper layers of the colony. Results demonstrate that FTIR microspectroscopy can be an useful tool for investigation of the spatial heterogeneity of cell growth within microbial macro-colonies. It is suggested that the method also can be adapted to the analysis of more complex multicellular communities, for example fruiting bodies, biofilms, or colonies growing under natural conditions.

Bacterial Physiological Phenomena↗

Optimization modeling of plant root architecture for water and phosphorus acquisition.

An optimization model is presented that examines the relationship between root architecture and multiple resource acquisition, specifically water and phosphorus in spatially heterogeneous environments. The basal root growth angle of an individual common bean plant, which determines the orientation and localization of the bulk of the root system, was modeled as the decision variable. The total payoff to the plant, the benefit obtained from water and phosphorus acquisition, minus the costs of spatial competition between roots, is given as a function of the (x,y) coordinates of the basal root in two-dimensional Cartesian space. We obtained a general solution and applied it to four unique environmental cases which are as follows: (1) the case of uniformly distributed water and phosphorus; (2) the case of localized shallow phosphorus; (3) the case of localized deep water; and (4) the case of shallow phosphorus and deep water. The general solution states that the optimal basal root growth angle will occur at the point where the total rate of change in the value of the resources acquired equals the total rate of change in cost that results from locating the root deeper in the soil. An optimizing plant locates its roots deeper in the soil profile until the marginal benefit exactly equals the marginal cost. The model predicts that the basal root angle of an optimizing plant will be shallower for Case 2 and deeper for Case 3, relative to the basal root angle obtained in the case of uniformly distributed water and phosphorus. The optimal basal root angle for Case 4 will depend on the marginal rate of substitution of water availability for phosphorus availability that occurs with depth. Empirical observations of bean root architecture in the greenhouse and in the field confirm model results and are discussed. In addition, the potential importance of phenotypic plasticity and phenotypic variation are discussed in relation to optimization of traits and adaptation to spatially heterogeneous environments.

Adaptation, Physiological↗

Molecular imaging of gene expression and efficacy following adenoviral-mediated brain tumor gene therapy.

Cancer gene therapy is an active area of research relying upon the transfer and subsequent expression of a therapeutic transgene into tumor cells in order to provide for therapeutic selectivity. Noninvasive assessment of therapeutic response and correlation of the location, magnitude, and duration of transgene expression in vivo would be particularly useful in the development of cancer gene therapy protocols by facilitating optimization of gene transfer protocols, vector development, and prodrug dosing schedules. In this study, we developed an adenoviral vector containing both the therapeutic transgene yeast cytosine deaminase (yCD) along with an optical reporter gene (luciferase). Following intratumoral injection of the vector into orthotopic 9 L gliomas, anatomical and diffusion-weighted MR images were obtained over time in order to provide for quantitative assessment of overall therapeutic efficacy and spatial heterogeneity of cell kill, respectively. In addition, bioluminescence images were acquired to assess the duration and magnitude of gene expression. MR images revealed significant reduction in tumor growth rates associated with yCD/5-fluorocytosine (5FC) gene therapy. Significant increases in mean tumor diffusion values were also observed during treatment with 5FC. Moreover, spatial heterogeneity in tumor diffusion changes were also observed revealing that diffusion magnetic resonance imaging could detect regional therapeutic effects due to the nonuniform delivery and/or expression of the therapeutic yCD transgene within the tumor mass. In addition, in vivo bioluminescence imaging detected luciferase gene expression, which was found to decrease over time during administration of the prodrug providing a noninvasive surrogate marker for monitoring gene expression. These results demonstrate the efficacy of the yCD/5FC strategy for the treatment of brain tumors and reveal the feasibility of using multimodality molecular and functional imaging for assessment of gene expression and therapeutic efficacy.

Adenoviridae↗

Spatial distribution of retinol-binding protein and retinyl palmitate hydrolase activity in normal and vitamin A-deficient rat liver.

A study was conducted to explore the spatial distribution within rat liver of two proteins importantly involved in retinoid metabolism in liver, namely, retinol-binding protein (RBP) and the enzyme retinyl palmitate hydrolase (RPH). The study was conducted with both vitamin A-sufficient (control) and vitamin A-deficient rats. Livers were carefully and reproducibly dissected into 11 sections each, and RBP levels and RPH activities were measured for each section homogenate. Both RBP and RPH activity displayed highly significant spatial heterogeneity in their distributions in liver. For control rats, the mean level of RBP was 39.0 micrograms/g wet weight, with a section-to-section variation of 14.5. For deficient rats, the corresponding RBP mean and variation values were 283 and 56 micrograms/g wet weight. For RPH, the mean level was 136 pmol free fatty acids (FFA) formed/(min X mg) with a section-to-section variation of 178. Both inspection of the data and analysis of variance indicated that this significant section-to-section variation (spatial heterogeneity) did not follow a consistent anatomic pattern from rat to rat. Thus, no one specific anatomic location in the liver was consistently high or low with regard to either RBP or RPH. Since the spatial distributions of both RBP and RPH activity did not follow a consistent anatomic pattern, it is not possible to obtain an accurate measure of the total liver levels for either parameter in a homogenate made from a small section. Finally, the patterns of distribution of RBP and RPH activity observed in the liver sections from both vitamin A-sufficient and deficient rats were not significantly correlated, either directly or inversely, as determined by chi-square analysis. Thus, RBP and RPH activity levels vary independently of each other in their heterogeneous anatomic distributions in rat liver.

Animals↗

Heterogeneity in spatial P-distribution and foraging capability by Zea mays: effects of patch size and barriers to restrict root proliferation within a patch.

BACKGROUND AND AIMS: Localized proliferation of roots in nutrient-enriched patches seems to be an adaptive response in many plants, but its function is still debatable. To understand the efficiency and limitation of foraging behaviour, the impact of patch size and the presence or absence of a barrier to root proliferation within phosphorus (P)-enriched patches was examined. METHODS: In pots filled with P-poor soil, six treatments of heterogeneous P supply were prepared: three patch sizes with or without a root barrier between patches. In addition, a homogeneous P supply treatment was also prepared. Irrespective of these treatments, each pot received the same total amount of P. Maize (Zea mays) was grown in each pot for 45 d in a greenhouse. KEY RESULTS: P content and biomass were greatest in plants grown in the largest patch due to successful root proliferation, and were higher in the presence of a root barrier. Interestingly, plants preferentially developed adventitious nodal roots projecting from the stem into the P-enriched soil, particularly in the largest patch with a root barrier. Removal of the barrier reduced the P-uptake capacity per unit root surface area or volume in P-enriched patches, revealing that the P-uptake capacity per root can be suppressed even in P-rich soil if other portions on the root axis encounter P-poor conditions. CONCLUSIONS: The results suggest that the efficiency of root morphological plasticity is largely determined by the size of the P-enriched patch. Furthermore, the results imply a novel aspect of P-uptake physiology that roots in heterogeneous P cannot demonstrate their potential capacity, as would be observed in roots encountering P continuously; this effect is probably mediated by an internal root factor.

Nitrogen↗