Agustín A. Grimoldi

Phosphorus Deficiency Decreases Cell Division and Elongation in Grass Leaves

Leaf growth in monocotyledons results from the flux of newly born cells out of the division zone and into the adjacent elongation-only zone, where cells reach their final length. We used a kinematic method to analyze the effect of phosphorus nutrition status on cell division and elongation parameters in the epidermis of Lolium perenne. Phosphorus deficiency reduced the leaf elongation rate by 39% due to decreases in the cell production rate (−19%) and final cell length (−20%). The former was solely due to a lower average cell division rate (0.028 versus 0.046 cell cell−1 h−1) and, thus, a lengthened average cell cycle duration (25 versus 15 h). The number of division cycles of the initial cell progeny (five to six) and, as a result, the number of meristematic cells (32–64) and division zone length were independent of phosphorus status. Accordingly, low-phosphorus cells maintained meristematic activity longer. Lack of effect of phosphorus deficiency on meristematic cell length implies that a lower division rate was matched to a lower elongation rate. Phosphorus deficiency did not affect the elongation-only zone length, thus leading to longer cell elongation duration (99 versus 75 h). However, the substantially reduced postmitotic average relative elongation rate (0.045 versus 0.064 mm mm−1 h−1) resulted in shorter mature cells. In summary, phosphorus deficiency did not affect the general controls of cell morphogenesis, but, by slowing down the rates of cell division and expansion, it slowed down its pace.

Escape from water or remain quiescent? Lotus tenuis changes its strategy depending on depth of submergence.

BACKGROUND AND AIMS Two main strategies that allow plants to cope with soil waterlogging or deeper submergence are: (1) escaping by means of upward shoot elongation or (2) remaining quiescent underwater. This study investigates these strategies in Lotus tenuis, a forage legume of increasing importance in areas prone to soil waterlogging, shallow submergence or complete submergence. METHODS Plants of L. tenuis were subjected for 30 d to well-drained (control), waterlogged (water-saturated soil), partially submerged (6 cm water depth) and completely submerged conditions. Plant responses assessed were tissue porosity, shoot number and length, biomass and utilization of water-soluble carbohydrates (WSCs) and starch in the crown. KEY RESULTS Lotus tenuis adjusted its strategy depending on the depth of submergence. Root growth of partially submerged plants ceased and carbon allocation prioritized shoot lengthening (32 cm vs. 24.5 cm under other treatments), without depleting carbohydrate reserves to sustain the faster growth. These plants also developed more shoot and root porosity. In contrast, completely submerged plants became quiescent, with no associated biomass accumulation, new shoot production or shoot elongation. In addition, tissue porosity was not enhanced. The survival of completely submerged plants is attributed to consumption of WSCs and starch reserves from crowns (concentrations 50-75 % less than in other treatments). CONCLUSIONS The forage legume L. tenuis has the flexibility either to escape from partial submergence by elongating its shoot more vigorously to avoid becoming totally submerged or to adopt a non-elongating quiescent strategy when completely immersed that is based on utilizing stored reserves. The possession of these alternative survival strategies helps to explain the success of L. tenuis in environments subjected to unpredictable flooding depths.

Different strategies of Lotus japonicus, L. corniculatus and L. tenuis to deal with complete submergence at seedling stage.

Two main strategies allow plants to deal with submergence: (i) escape from below water by means of shoot elongation, or (ii) remaining quiescent under the water until water subsides and then resume growth. We investigated these strategies in seedlings of Lotus japonicus, L. corniculatus and L. tenuis subjected to control and submergence for 12 days, with a subsequent 30-day recovery period. All three species survived submergence but used different strategies. Submerged seedlings of L. japonicus exhibited an escape strategy (emerging from water) as a result of preferential carbon allocation towards shoot mass and lengthening, in detriment to root growth. In contrast, seedlings of L. corniculatus and L. tenuis became quiescent, with no biomass accumulation, no new unfolding of leaves and no shoot elongation. Upon de-submergence, seedlings of L. japonicus had the lowest recovery growth (a biomass and shoot height 58% and 40% less than controls, respectively), L. corniculatus was intermediate and L. tenuis showed the greatest recovery growth. Previously submerged seedlings of L. tenuis did not differ from their controls, either in final shoot biomass or shoot height. Thus, for the studied species, quiescence appears to be an adequate strategy for tolerance of short-term (i.e., 12 days) complete submergence, being consistent with field observations of L. tenuis colonisation of flood-prone environments.

Flooding Effects on Plants Recovering from Defoliation in Paspalum dilatatum and Lotus tenuis

BACKGROUND AND AIMS Flooding and grazing are major disturbances that simultaneously affect plant performance in many humid grassland ecosystems. The effects of flooding on plant recovery from defoliation were studied in two species: the grass Paspalum dilatatum, regrowing primarily from current assimilation; and the legume, Lotus tenuis, which can use crown reserves during regrowth. METHODS Plants of both species were subjected to intense defoliation in combination with 15 d of flooding at 6 cm water depth. Plant recovery was evaluated during a subsequent 30-d growth period under well-watered conditions. Plant responses in tissue porosity, height, tiller or shoot number and biomass of the different organs were assessed. KEY RESULTS Flooding increased porosity in both P. dilatatum and L. tenuis, as expected in flood-tolerant species. In P. dilatatum, defoliation of flooded plants induced a reduction in plant height, thus encouraging the prostrated-growth response typical of defoliated plants rather than the restoration of contact with atmospheric oxygen, and most tillers remained submerged until the end of the flooding period. In contrast, in L. tenuis, plant height was not reduced when defoliated and flooded, a high proportion of shoots being presented emerging above water (72 %). In consequence, flooding plus defoliation did not depress plant recovery from defoliation in the legume species, which showed high sprouting and use of crown biomass during regrowth, whereas in the grass species it negatively affected plant recovery, achieving 32 % lower biomass than plants subjected to flooding or defoliation as single treatments. CONCLUSIONS The interactive effect of flooding and defoliation determines a reduction in the regrowth of P. dilatatum that was not detected in L. tenuis. In the legume, the use of crown reserves seems to be a key factor in plant recovery from defoliation under flooding conditions.

CONSTITUTIVE AND PLASTIC ROOT TRAITS AND THEIR ROLE IN DIFFERENTIAL TOLERANCE TO SOIL FLOODING AMONG COEXISTING SPECIES OF A LOWLAND GRASSLAND

Natural flooding is a major component of the disturbance regime in many grassland ecosystems. The objective of this study was to analyze the relationship among constitutive and plastic root traits and tolerance to flooding in coexisting perennial species of the flooding pampa grasslands (Argentina). A mesocosm experiment was designed for five native species (Paspalidium paludivagum, Paspalum dilatatum, Bothriochloa laguroides, Eryngium ebracteatum, and Eclipta bellidioides) and two exotic ones (Mentha pulegium and Plantago lanceolata). Across species, constitutive root porosity was positively correlated with the tolerance to soil flooding. Moreover, the generation of additional aerenchyma was larger in species with intermediate values of constitutive root porosity and lower in species with low or high constitutive root porosity. This differential increase in the root porosity of each species, combined with the values of constitutive root porosity, resulted in a stronger correlation between final root porosity and tolerance under flooding conditions. Native grasses increased the proportion of root aerenchyma, showing a small change in the number of lysed cells but a significant increase in the cortex proportion and diameter of roots. Exotic dicots generated lysigenous aerenchyma throughout their cortex; in contrast, native dicot species maintained the cell layers adjacent to the stele. A lag in the development of secondary growth during flooding was detected in both groups of dicots, a response that was particularly evident in the exotic species, contrasting with their prominent growth under nonflooded conditions. In general, our results indicate that constitutive and plastic root traits are very significant in terms of the effects of periodic flooding on the abundance of coexisting species of the flooding pampa grasslands.

The effects of submergence on anatomical, morphological and biomass allocation responses of tropical grasses Chloris gayana and Panicum coloratum at seedling stage

Abstract. Submergence is a major factor affecting seedling recruitment in lowland grassland ecosystems. Our aim was to evaluate the tolerance to increasing flooding intensity of the seedlings of tropical grasses Chloris gayana K. and Panicum coloratum L., whose use as a forage species is increasing in humid grasslands. For this purpose, 2-week-old seedlings of C. gayana and P. coloratum were subjected to control, partial submergence (PS) and complete submergence (CS) in clear water for 14 days and allowed to grow for a subsequent 12-day period to assess their recovery. The following responses were assessed: generation of root aerenchyma, morphological changes and emergence from water, biomass allocation in relation to plant size, and biomass accumulation. Results showed that constitutive root aerenchyma was high in both species. Under PS and CS, root aerenchyma increased by up to 50–55% in C. gayana and up to 40–48% in P. coloratum. Under PS, the increase in seedling height for both species was the same as for controls. Under CS, C. gayana further increased its height and emerged more quickly from water; P. coloratum was not able to increase its height, and therefore the seedlings always remained underwater. The escape-from-water response of C. gayana was associated with preferential biomass allocation towards shoots and with a marked lengthening of leaf blades. By contrast, there was no change in allocation in P. coloratum, and its leaves were shorter under CS. The final biomass of C. gayana under CS was similar to that under PS, and equivalent to 54% of its controls. In P. coloratum, biomass under PS and CS were 64 and 21% of its controls (respectively), which indicates that injury caused by CS persisted during the post-submergence period. In conclusion, both species are tolerant to PS at the seedling stage. However, when flood depth increases by submerging the seedlings, C. gayana is able to escape from water while P. coloratum is not, thus strongly affecting its recovery. Therefore, C. gayana appears to be a more promising species for cultivation in lowland grasslands prone to flooding of unpredictable intensity.

Soil water regime of grassland communities along subtle topographic gradient in the Flooding Pampa (Argentina).

Di Bella C.E., Striker G.G., Loreti J., Cosentino D.J., Grimoldi A.A. (2016): Soil water regime of grassland communities along subtle topographic gradient in the Flooding Pampa (Argentina). Soil & Water Res., 11: 90–96. Three plant communities positioned along a subtle topographic gradient, referred to as upland, intermediate, and lowland positions, characterize the landscape of the Flooding Pampa grasslands of Argentina. Although it is believed that the structure and functioning of the plant communities at each position are in close relationship with their hydric regime, this has never been quantified. More than 800 measurements of soil water content during four years, along with soil water retention curves, and physical and chemical parameters of soils were assessed at each position. Results showed that water availability during the year varied among the positions in accordance with differences in hydrological balance and soil water retention capacity of each of them. Water retention increased in relation to clay and organic matter content from the upland to the lowland position. The upland position, with more soil sand content, registered severe drought events during late spring and summer, without flooding periods in any season. The intermediate and lowland positions, with more soil clay content, remained flooded for several weeks during winter and spring, and they manifested less severe summer droughts than the upland position. Moreover, the lowland position was more hydromorphic than the intermediate one. These spatial and temporal variations of water regime and soil parameters characterizing the upland, intermediate, and lowland positions concur with different plant communities associated with each of them.

Differential growth of Spartina densiflora populations under saline flooding is related to adventitious root formation and innate root ion regulation

Global change anticipates scenarios of sea level rise that would provoke long lasting floods, especially in lowland areas of salt marshes. Our aim was to evaluate the morpho-physiological adjustment ability to deal with continuous saline flooding of Spartina densiflora Brogn. plants from lowlands and uplands along a subtle topographical gradient (0.2m differential altitude). Plants from both origins were subjected to continuous saline flooding (300mM NaCl) for 35 days. Responses associated to adventitious rooting, aerenchyma formation, concentration of Na+, K+ and Cl- in roots and shoots tissues, tillering and growth were assessed. Root responses differentiated populations given that lowland plants showed higher ability for adventitious root formation and innate superior root ion regulation than upland plants. High constitutive K+ concentration plus high Na+ exclusion in root tissues led to significant low values of Na+:K+ ratios in lowland plants. Better root functioning was, in turn, related with more consistent shoot performance as lowland plants maintained plant tiller number and shoot relative growth rate unaltered while upland plants decreased both parameters by 35 and 18%, respectively, when in saline flooding. The superior performance of lowland plants indicates that locally adapted populations can be promoted in salt marsh habitats with subtle differences at topographic level.

Ability to recover overrides the negative effects of flooding on growth of tropical grasses Chloris gayana and Panicum coloratum

Abstract. This study assessed the flooding tolerance of the tropical grasses Chloris gayana Kunth and Panicum coloratum L. at different times of the year: (i) late winter flooding for 50 days (WF), (ii) early spring flooding (SF) for 20 days, and (iii) long-term flooding covering both periods (WF + SF, 70 days). A growth period under well-watered conditions was allowed after each flooding event to assess recovery of plant species. Plants were harvested after each flooding event and at the end of the recovery period. Panicum coloratum had higher tolerance to WF than C. gayana. Treatment WF did not affect biomass in P. coloratum, whereas it reduced biomass of flooded plants by 38% in C. gayana. Treatment SF did not differentiate the species for tolerance; both registered moderate reduction in their growth (20–30%). Under WF + SF, C. gayana showed additional reduction in its growth over that observed when subjected separately to either WF or SF, whereas P. coloratum did not. Both species displayed remarkably fast recovery from flooding when temperatures rose during early summer, attaining biomass equivalent to that of non-flooded plants 1 month after water subsided. Therefore, although P. coloratum appears slightly more tolerant during flooding than C. gayana, both species are promising for introduction in temperate lowland grasslands.

Growth during recovery evidences the waterlogging tolerance of forage grasses

Abstract. Waterlogging is a stress of increasing importance for pastures as a consequence of global climate change. We evaluated the impact of waterlogging on four forage grasses with alleged differential tolerance, emphasising not only responses during the stress but also their reported ability to recover from it. To do this, 42-day plants of Dactylis glomerata, Bromus catharticus, Festuca arundinacea and Phalaris aquatica were subjected to 15-day waterlogging, followed by a subsequent 15-day recovery period. Shoot and root growth (i.e. RGR) during both periods, in addition to net photosynthesis and stomatal conductance rates during waterlogging were assessed. Sensitivity exhibited by D. glomerata and B. catharticus during waterlogging was related to growth arrest of roots – but not of shoots – along with a progressive fall in stomatal conductance and net photosynthesis. The injury during waterlogging preceded a negligible growth of shoots and roots, only evident during recovery in both species. By contrast, P. aquatica exhibited unaltered root RGR and promoted shoot RGR with no impact on leaf gas exchange during waterlogging; whereas F. arundinacea showed intermediate tolerance as root RGR was reduced during waterlogging, with stomatal conductance, net photosynthesis and shoot RGR remaining unaffected. These latter two species fully regained shoot and root RGR during recovery. So, P. aquatica and F. arundinacea seem more suitable for prone-to-flood lowlands, whereas to be conclusive about waterlogging tolerance, it is necessary to examine plant recovery as shown in D. glomerata and B. catharticus.