Plants display an array of classical strategies to maximize phosphate (Pi) acquisition from sparingly soluble P sources. Acclimation to Pi-stress via elegant Pi-starvation induced (PSI) adjustments would reduce our current overreliance on expensive, polluting and non-renewable Pi-fertilizers. Nevertheless, differences in the ability of various species to solubilize sparingly soluble P-sources have been often evidenced; inter-cultivar variations are scarcely documented. Brassica is known as an effective, non-mycorrhizal user of sparingly soluble P-sources. Various growth parameters and biomass accumulation by genetically diverse Brassica cultivars were determined in four experiments using hydroponics and quartz sand culture media. Role of PSI root mediated pH changes, organic anions (OAs) exudation and altered root architecture in mobilization and acquisition of sparingly soluble P-forms [Jordan rock-P (RP) at 2 g l )1 and Ca 3 (PO 4 ) 2 (TCP) at 0.2 g l )1 respectively] was investigated. Cultivars showed considerable genetic variations in biomass accumulation, various growth parameters and root-shoot ratio. Concentration and total uptake of P, specific absorption rate of P, P-transport rate and P-utilization efficiency (PUE) were also significantly (P < 0.001) different for various cultivars and their dry matter was significantly correlated with P-uptake [r = 0.94** (significant at 1% level)]. P-tolerant cultivars showed substantial decrease in solution media pH because of H + efflux and exuded more carboxylates than low P-sensitive cultivars under P-starvation. P-uptake by cultivars increased linearly with decreasing pH. The amount and types of OAs exuded from the roots of P-starved plants differed from those of plants grown under P-sufficient environment. In split pot study, with TCP and RP supplied spatially separated from other nutrients, efficient cultivars were still able to mobilize RP and TCP more efficiently than inefficient cultivars. In rhizobox study, the elongation rates of primary roots decreased but the elongation rates of the branched zones of primary roots and the length of lateral roots increased under P-starvation. Tested cultivars showed genetic diversity in accessing, mobilization, acquisition and utilization of Pi from sparingly soluble P forms. An arrange marriage of plant traits can explain cultivar's access to different forms of sparingly soluble P, and in addition to altered lateral root topology and enhanced P-uptake and PUE, enhanced H + efflux and OAs exudation are key factors in Pi scavenging from extra cellular sparingly soluble P-forms.
Translocation of absorbed phosphorus (P) from metabolically inactive sites to active sites in plants growing under P deprivation may increase its P utilization efficiency (PUE). Acclimation to phosphate (Pi) starvation may be caused by a differential storage pool of vacuolar P, its release, and the intensity of re-translocation of absorbed P as P starvation inducible environmental cues (PSIEC) from ambient environment. Biomass assay and three P forms, namely inorganic (Pi), organic (Po), and acid-soluble total (Ptas) were estimated in Brassica cultivars exposed to 10 d P deprivation in the culture media. Considering that -delta Pi/delta t denotes the rate of Pi release, Pi release velocity (RSPi) was determined as the tangent to the equations obtained for Pi f(t) at the mean point in the period of greatest Pi decrease, whereas the inverse of the RSPi was an estimate of the internal Pi buffering capacity (IBCPi). Inter cultivar variations in size of the non-metabolic Pi pool, RSPi, re-translocation of Pi from less to more active metabolic sites, and preferential Pi source and sink compartments were evaluated under P starvation. The cultivar 'Brown Raya' showed the highest Pi storage ability under adequate external P supply, and a more intensive release than 'Rain Bow' and 'Dunkled' under P stress. Cultivar 'B.S.A' was inferior to 'Con-1' in its ability to store and use Pi. Roots and upper leaves were the main sink of Pi stored in the lower and middle leaves of all cultivars and showed lower IBCPi and larger RSPi values than lower and middle leaves. In another trial, six cultivars were exposed to P-free nutrition for 29 d after initial feeding on optimum nutrition for 15 d. With variable magnitude, all of the cultivars re-translocated P from the above ground parts to their roots under P starvation, and [P] at 44 d after transplanting was higher in developing leaves compared with developed leaves. Under P deprivation, translocation of absorbed P from metabolically inactive to active sites may have helped the tolerant cultivars to establish a better rooting system, which provided a basis for tolerance against P starvation and increased PUE. A better understanding of the extent to which changes in the flux of P absorption and re-translocation under PSIEC will help to scavenge Pi from bound P reserves and will bring more sparingly soluble P into cropping systems and obtain capitalization of P reserves.
In calcareous soils, phosphorus (P) retention and immobilization take place due to precipitation and adsorption. Since soil pH is considered a major soil variable affecting the P sorption, an acidic P fertilizer could result in low P adsorption compared to alkaline one. Therefore, P adsorption from DAP and phosphoric acid (PA) required to produce desired soil solution P concentration was estimated using Freundlich sorption isotherms. Two soils from Faisalabad and T. T. Singh districts were spiked with 0, 10, and 20 % CaCO3 for 15 days. Freundlich adsorption isotherms (P = aC b/a) were constructed, and theoretical doses of PA and DAP to develop a desired soil solution P level (i.e., 0.20 mg L−1) were calculated. It was observed that P adsorption in soil increased with CaCO3. Moreover, at all the levels of CaCO3, P adsorption from PA was lower compared to that from DAP in both the soils. Consequently, lesser quantity of PA was required to produce desired solution P, 0.2 mg L−1, compared to DAP. However, extrapolating the developed relationship between soil CaCO3 contents and quantity of fertilizer to other similar textured soils needs confirmation.
Plants grown in highly weathered or highly alkaline calcareous soils often experience phosphorus (P) stress but never a P-free environment. Thus, applications of mineral P fertilizers are often required to achieve maximum yield, but recovery of applied P fertilizers is notoriously low. Phosphorus deprivation elicits a complex array of morphological, physiological, and biochemical adaptations among plant species and genotypes to enhance P acquisition and utilization efficiency. Ten Brassica cultivars were grown hydroponically to investigate their relative efficiency to utilize deficiently (20-mM) and adequately (200-mM) supplied P, using Johnson's modified solution. Cultivars differed significantly (P , 0.001) in biomass accumulation. Orthophosphate concentration and uptake in shoot and root, absolute and relative growth rate, and P-utilization efficiency (PUE) were also significantly different among various Brassica cultivars. Root-shoot ratio and specific absorption rate were substantially increased in plants subjected to low P supply. Shoot and root dry-matter yield as well as total biomass production correlated significantly (P , 0.01) with their total P uptake and PUE. Cultivars, which were efficient in P utilization, were also efficient accumulators of biomass under adequate as well as deficient levels of P supply. As part of the study, kinetic parameters of P uptake were evaluated for six contrasting Brassica cultivars in PUE, grown in nutrient solution. The kinetic parameters related to P influx were maximal transport rate (Vmax), the Michaelis-Menten constant (Km), and the external concentration when net uptake is zero (Cmin). Lower Km and Cmin values were indicative of P-uptake ability of the cultivars, evidencing their adaptability to P-stress conditions. In another experiment, six cultivars were exposed to no P nutrition for 27 days after initial feeding on optimum nutrition for 14 days. All the cultivars retranslocated P from aboveground parts to their roots during growth in P-free conditions, the magnitude of which was variable in different cultivars. Phosphorus concentration at 41 days after transplanting was higher in developing leaves than developed leaves. Translocation of absorbed P from metabolically inactive sites to active sites in plants growing under P-stress conditions may have helped the tolerant cultivars to establish a better rooting system, which provided basis for tolerance against P-deficiency stress and increased PUE.
Non-mycorrhizal Brassica does not produce specialized root structures such as cluster or dauciform roots but is an effective user of P compared with other crops. In addition to P-uptake, utilization and remobilization activity, acquisition of orthophosphate (Pi) from extracellular sparingly P-sources or unavailable bound P-forms can be enhanced by biochemical rescue mechanisms such copious H(+)-efflux and/or carboxylates exudation into rhizosphere by roots via plasmalemma H(+) ATPase and anion channels triggered by P-starvation. To visualize the dissolution of sparingly soluble Ca-phosphate (Ca-P), newly formed Ca-P was suspended in agar containing other essential nutrients. With NH(4)(+) applied as the N source, the precipitate dissolved in the root vicinity can be ascribed to rhizosphere acidification, whereas no dissolution occurred with nitrate nutrition. To observe in situ rhizospheric pH changes, images were recorded after embedding the roots in agar containing bromocresol purple as a pH indicator. P-tolerant cultivar showed a greater decrease in pH than the sensitive cultivar in the culture media (the appearance of typical patterns of various colors of pH indicator in the root vicinity), and at stress P-level this acidification was more prominent. In experiment 2, low P-tolerant class-I cultivars (Oscar and Con-II) showed a greater decrease in solution media pH than low P-sensitive class-II (Gold Rush and RL-18) cultivars, and P-contents of the cultivars was inversely related to decrease in culture media pH. To elucidate P-stress-induced remodeling and redesigning in a root architectural system, cultivars were grown in rhizoboxes in experiment 3. The elongation rates of primary roots increased as P-supply increased, but the elongation rates of the branched zones of primary roots decreased. The length of the lateral roots and topological index values increased when cultivars were exposed to a P-stress environment. To elucidate Pi-uptake kinetics, parameters related to P influx: maximal transport rate (V(max)), the Michaelis-Menten constant (K(m)), and the external concentration when net uptake is zero (C(min)) were tested in experiment 4. Lower K(m) and C(min) values were better indicative of the P-uptake ability of the class-I cultivars, evidencing their adaptability to P-starved environmental cues. In experiment 5, class-I cultivars exuded two- to threefold more carboxylates than class-II cultivars under the P-stress environment. The amount and types of carboxylates exuded from the roots of P-starved plants differed from those of plants grown under P-sufficient conditions. Nevertheless, the exudation rate of both class-I and class-II cultivars decreased with time, and the highest exudation rate was found after the first 4 h of carboxylates collection. Higher P uptake by class-I cultivars was significantly related to the drop in root medium pH, which can be ascribed to H(+)-efflux from the roots supplied with sparingly soluble rock-P and Ca(3)(PO(4))(2). These classical rescue strategies provided the basis...
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