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Plant Water Deficits, Osmotic Properties, and Hydraulic Resistances of Hawaiian Dubautia Species from Adjacent Bog and Wet-Forest Habitats!

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Plant Water Deficits, Osmotic Properties, and Hydraulic Resistances of Hawaiian Dubautia Species from Adjacent Bog and Wet-Forest Habitats! Pacific Science (1990), vol. 44, no. 4: 449-455 © 1990 by University of Hawaii Press. All rights reserved Plant Water Deficits, Osmotic Properties, and Hydraulic Resistances of Hawaiian Dubautia Species from Adjacent Bog and Wet-Forest Habitats! JOAN E. CANFIELD 2 ABSTRACT: Functional responses oftwo closely related Dubautia species from a mosaic of Hawaiian bogs and wet forest were compared to help explain their differential distributions. Dubautia paleata is largely restricted to saturated bogs, while D. raillardioides is restricted to the surrounding, better-drained wet forest. Minimum diurnal tissue water potentials of D. paleata are significantly lower than those of D. raillardioides, despite the moister condition of bog soil. The tissue osmotic potential at full hydration (nJ of D. paleata is significantly lower than that of D. raillardioides. As a result , the tissue water potential at which turgor reaches zero for D. paleata is significantly lower than that of D. raillardioides. Dubautia paleata is thus able to maintain positive turgor to lower water poten­ tials than D. raillardioides. Lack ofa lowered n, in D. raillardioides may therefore contribute to exclusion of that species from the bog habitat. Preliminary data suggest a significantly greater hydraulic resistance for D. paleata than for D. raillardioides, probably due to higher root resistance caused by the reduced condition of the waterlogged bog substrate. The difference in hydraulic resis­ tance could help account for the contrasting water deficits of the two species. As ONE OF THE WETTEST terrestrial habitats, creating sucstantial hydraulic resistance in bogs would not be expected to cause substan­ bog plants (Marchand 1975, Bradbury and tial tissue water deficits in plants. However, Grace 1983). moderately low tissue water potentials (-1.5 One physiological mechanism capable of to - 2 MPa) have been reported for temperate buffering the effect of tissue water deficits and subarctic bog species (Small 1972, is a decrease in tissue osmotic potential. A Johansson and Linder 1975, Marchand 1975). lowered osmotic potential enables turgor to Although these minima are not nearly as low be maintained to lower tissue water contents as those reported for xerophytes, they never­ (Tyree and Jarvis 1982). Stomatal opening theless may affect metabolic processes ad­ and photosynthetic carbon assimilation are versely (Hsiao et al. 1976). among the physiological processes adversely One probable cause oflow water potentials affected by water deficits. Whether moderate in bog plants is the saturated soil. Hypoxia water deficits are transduced into metabolic inhibits root respiration, reducing the viable changes through a reduction in turgor pressure root mass (Bannister 1964, Cook et al. 1980). is still a matter of debate (Hsiao et al. 1976, Appreciable root resistance can thus develop , Turner and Jones 1980, Schulze 1986). It has been argued that the ability ofplants to main- tain turgor at lower tissue water contents may promote their growth and survival under 1 This research was supported by National Science Foundation Grant DEB79-I0993 to Dieter Mueller­ moisture-limited conditions (Turner and Jones Domboi s and by the Hawaii Evolutiona ry Biology Pro­ 1980, Bradford and Hsiao 1982).Low osmotic gram. Manuscript accepted 15January 1990. potentials (e.g., -3 MPa) have been reported 2 Department of Botany, University of Hawaii at for bog plants (Harris 1934, Ingram 1983). Manoa, Honolulu, Hawaii 96822. Present address: U.S. Fish and Wildlife Service, Pacific Islands Office, Office of This paper addresses the water status ofbog Fish and Wildlife Enhancement, P.O. Box 50167, Hono­ plants by examining two closely related spe­ lulu, Hawaii 96850. cies of Dubautia (Asteraceae) from adjacent 449 450 PACIFIC SCIENCE, Volume 44, October 1990 bog and wet-forest habitats in Hawaii. The ~ - 4 I extent to which these plants develop tissue 0 water deficits is described first. The osmotic n, 6 properties of the species are discussed, along -3 0 with the potential effect of those properties on + Dubautia paleata turgor maintenance. Finally, limited hydrau­ c ......Q) lic resistance data are related to the contrast­ 0 ing water deficits of the two species. Q. - 2 "- ......Q) 0 ~ -1 MATERIALS AND METHODS Q) .... lf) "- Q) Dubautia paleata Gray and D. raillardioides > .................................... Hbd. are endemic to the Hawaiian island of c 0 Kauai. They are most common in the central 1.0 0.8 0.6 0.4 0.2 0.0 upland portion ofKauai known as the Alakai Relative water content Swamp, a dissected plateau of wet forest and open bogs within the montane rainforest FIGURE I. The reciprocal of tissue water potential as a zone. Located at 1200- 1600 m elevation, the function of tissue relative water content for a branch of Alakai plateau receives abundant orographic D. paleata . One additional point extends the curve higher on the ordinate than shown here. The correlation coeffi­ precipitation brought by the prevailing north­ cient for the linear region of the curve (defined by seven east tradewinds. The average annual rainfall data points) is 0.999. ranges from 2500 to 11,500 mm and is not markedly seasonal (State of Hawaii 1982). The study site, Aipo Iki bog along the Tissue osmotic properties were determined Alakai Swamp Trail (1222 m elevation; 3200 by the pressure chamber technique (Tyree mm annual rainfall) , is a typical open to and Jarvis 1982, Robichaux 1984). Sets of shrubby, nearly flat, poorly drained bog sur­ branches were collected during the summer rounded by low-stature wet forest near better­ and winter of 1983and the spring and summer drained slopes. In both habitats Metrosideros of 1984. After being recut under water in the polymorpha (Myrtaceae) is dominant. The laboratory, the branches were stored over­ profusion of epiphytic ferns and bryophytes night in distilled water to restore turgor. Re­ attests to the prevalence of rain and fog at the peated measurements of weight and water site. Highly acidic, saturated peat forms the potential were made as the branches dried out, bog substrate. In the adjacent forest the soil following the procedure of Robichaux (1984). water content averages half that of the bog, Dry weight ofthe branches was obtained after and the rhizosphere is typically three times as oven drying for 24 hr at 70-80° C. Weights deep as in the bog (Canfield 1986). were measured with a Sartorius GmbH model Dubautiapaleata is a small shrub, averaging 1212MP digital balance. 0.5 m in height, and is primarily restricted The relative water content of each branch to the bog. Dubautia raillardioides is a large, was calculated as described by Robichaux sprawling shrub, often reaching a height of 2 (1984). The inverse of tissue water potential m, and is confined to the understory ofthe wet was then plotted as a function of relative forest. Because the boundary between the two water content (Figure I). Extrapolation of the habitats is rather sharp, individuals ofthe two linear part of the resulting curve to the ordi­ species can be found growing within 20 m of nate gives the reciprocal ofthe osmotic poten­ each other. tial at full hydration (Tyree and Jarvis 1982). Diurnal tissue water potentials of the two The difference between the values of water species were determined during the summer of potential (on the curve) and osmotic potential 1983 and spring, summer, and winter of 1984 (on the extrapolated line) for a given value of with a PMS Instrument Co. model 1000 pres­ relative water content is equivalent to the sure chamber. tissue turgor pressure (Tyree and Jarvis 1982). Water Deficits in Hawaiian Dubautia-CANFIELD 451 Hydraulic resistances to liquid water flux in 0.01''''''"'!'"'----------__, the soil-root-shoot continuum were calcu­ lated for five to six branches of each species '0' on 16 July 1985 with the following formula: c, -0.5 R = (l/Jbran ch -l/Jsoil)/E, where R is hydraulic .6 resistance, l/Jbranch and l/Jsoil are tissue and soil :2..... water potentials, respectively, and E is trans­ 55 -1.0 piration (Cowan 1977, Calkin and Pearcy 15 1984). This formula makes the assumption a. I- that E can be used as an estimate ofthe liquid ~ water flux supplied per unit area of leaf by ~ -1.5 the soil-root-shoot continuum (Calkin and o D. paleata • D. raillardioides Pearcy 1984).l/Jsoil was determined with a Soil­ moisture Equipment Co. model 1500 ceramic plate extractor. Values of l/Jsoil were> -0.01 Weather MPa for both species. ~ i V(l//221 E was calculated according to Cowan 7 11 15 19 (1977) and Hall and Schulze (1980) except for Local time (hr) the boundary layer resistance to water vapor FIGURE 2. Diurnal tissue water potentials of D. paleata (rb ) . rb was calculated as [O.4(d/u)-o.S] /D, from Aipo Iki bog and D . raillardioides from the adjacent where d is leaf width , u is wind speed, and D wet forest on 25 March 1984. Vertical lines show the range is the diffusion coefficient of water vapor of variation for duplicates, where not eclipsed by the (Campbell 1977, Nobel 1983). Wind speeds symbol. Weather conditions indicated at bottom are were measured with a Weather Measure partl y cloudy (shaded) or clear and sunny (not shaded). Corp. model Wl31 anemometer. For the calculation ofE, leaf resistances to period, the mean value of l/Jmin for D. paleata water vapor were measured with a LI-Cor Inc. (- 1.07 ± 0.13 MPa) was significantly lower model LI-1600 steady-state porometer. Leaf (P < 0.001) than that of D. raillardioides temperatures were measured with fine-wire ( - 0.47 ± 0.17 MPa). Maximum tissue water (0.076 mm) copper-constantan thermo­ potentials (l/Jmax) of D. paleata were the same couples appressed to the abaxial leaf surfaces as or slightly higher than those of D.
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