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- Coordinated Tree Responses to Drought -Vulnerability and Sustainable Production: Hypotheses on Arid Ecosystem Adjustments to Limitations in Water Resources (2004)
- Field and controlled greenhouse experiments were carried out to investigate tree responses to declining soil water content. Field experiments were conducted on naturally growing trees of Acacia tortilis and A. xanthophloea in the savanna region of Kenya and Quercus suber in the Mediterranean region of Portugal. The selected field sites were regions that experience regular drought periods during the year. Greenhouse experiments constituted two watering regimes. Seedlings of A. tortilis and A. xanthophloea grown from seeds initially obtained from the Kenya field site were raised and arranged on a greenhouse bench into two groups per species. The first set of plants were watered every other day (controls) while the second set were watered every seven days (water stress treatments). Field measurements included weather parameters, soil and plant water status, growth, sap flux density, leaf transpiration and stomatal conductance, tissue water relations and isotope labeling. Similar measurements were conducted on plants growing in the greenhouse. Also examined in the greenhouse were root biomass, root structure as well as whole plant biomass accumulation. A second set of experiments was carried out in the greenhouse by subjecting plants initially stressed and non-stressed to severe water stress by withholding water until plants were wilted overnight. The wilted plants were then re-watered regularly and their recovery after stress alleviation was monitored. Declining soil water content significantly affected plant water status in all the trees studied. Lowest psi pd recorded during the study period occurred in the month of June and were –2.0 and –1.1 MPa for A. xanthophloea and A. tortilis respectively. The same species subjected to repeated water stress in the greenhouse attained mean minimum psi pd of –2.4 and –1.2 MPa for A. xanthophloea and A. tortilis respectively at the end of the drying cycle. Mean minimum psi pd recorded for Q. suber during summer was –1.8 MPa and occurred in September. There were however, significant differences among trees. Decline in psi associated with increasing soil drought led to decline in leaf initiation and leaf expansion and both processes ceased at higher water stress levels. For the Acacia species, even leaf shedding occurred at higher stress levels. There was also a decline in stomatal conductance (gs) during water stress, leading to decrease in transpiration rates (E). Maximum stomatal conductance of 340 mmol m-2 s-1 were observed during rainy seasons for the Acacia trees while mean maximum values of 300 mmol m-2 s-1 were recorded for Q. suber when soil moisture conditions were favorable. Stomatal conductance declined by 31%, 67% and 67% in A. tortilis and A. xanthophloea in the savanna and Q. suber in the Mediterranean regions respectively. Daily tree water use (Qtree) as well as leaf transpiration reflected changes in psi and gs. Root to leaf hydraulic conductance equally declined with increasing soil drought. Q. suber trees adjusted osmotically by a magnitude of 0.7 MPa, while bulk modulus of elasticity (epsilon) increased by a magnitude of 17 MPa. Osmotic adjustment of 0.48 MPa was observed in greenhouse plants of A. tortilis while epsilon declined by a magnitude of 7 MPa in A. xanthophloea. A. tortilis plants in the greenhouse showed increased absolute root growth, root depth and root:shoot (r:s) ratio. The dimorphic rooting pattern in Q. suber resulted into hydraulic lift and this could as well occur in A. tortilis because of similarity in their rooting patterns. Most plant responses were reactionary and were aimed at enhancing soil water uptake and reducing transpiration water loss when soil water content was declining. Similar responses were observed for both greenhouse and naturally growing field plants of the same species. Decline in leaf initiation and leaf expansion as well as leaf senescence reduced tree crown size hence potential tree transpiration. This however, had negative impact on plant productivity. Increased root growth as well as osmotic adjustment increased tree water uptake from the soil. The balance between root water uptake and leaf transpiration through growth and stomatal regulation was aimed at protecting xylem integrity. The overall results showed that soil characteristics, root activities and root distribution patterns are the main factors determining tree functioning and productivity in drylands, while the coordinated interaction between the aboveground shoot and belowground root activities ensures survival during drought. Maintained production and survival will ensure distribution and success in the arid environments. Repeated water stress imparted water stress resistance qualities on seedlings enabling them to survive longer during severe stress. The study emphasizes the role of soil resource base as well as species interactions in the functioning and balance of dryland ecosystems.