Baobab trees (Adansonia) in Madagascar use stored water to ﬂush new leaves but not to support stomatal opening before the rainy season
Blackwell Publishing Ltd
Saharah Moon Chapotin1, Juvet H. Razanameharizaka2 and N. Michele Holbrook1
Department of Organismic and Evolutionary Biology, Harvard University, Cambridge, MA 02138, USA; 2Département de Biologie et Ecologie Végétales,
Université d’Antananarivo, Antananarivo (101), Madagascar
Author for correspondence: Saharah Moon Chapotin Tel: +1 617 496 3580 Fax: +1 617 496 5854 Email: email@example.com Received: 11 August 2005 Accepted: 10 October 2005
• Baobab trees (Adansonia, Bombacaceae) are widely thought to store water in their stems for use when water availability is low. We tested this hypothesis by assessing the role of stored water during the dry season in three baobab species in Madagascar. • In the dry season, leaves are present only during and after leaf ﬂush. We quantiﬁed the relative contributions of stem and soil water during this period through measures of stem water content, sap ﬂow and stomatal conductance. • Rates of sap ﬂow at the base of the trunk were near zero, indicating that leaf ﬂushing was almost entirely dependent on stem water. Stem water content declined by up to 12% during this period, yet stomatal conductance and branch sap ﬂow rates remained very low. • Stem water reserves were used to support new leaf growth and cuticular transpiration, but not to support stomatal opening before the rainy season. Stomatal opening coincided with the onset of sap ﬂow at the base of the trunk and occurred only after signiﬁcant rainfall. Key words: Adansonia, Bombacaceae, leaf ﬂushing, phenology, sap ﬂow, stomatal conductance, tropical dry forest, water storage. New Phytologist (2006) 169: 549–559 © The Authors (2005). Journal compilation © New Phytologist (2005) doi: 10.1111/j.1469-8137.2005.01618.x
Stem-succulent trees are prominent in arid tropical ecosystems around the world, and baobab trees (Adansonia L.; Bombacaceae) are among the best-known examples. Because of the high water content and large volume of their stems and branches, and their occurrence in seasonally dry environments, it has long been assumed that baobab trees depend on stored water during periods of low water availability, and replenish the stored water during periods of rainfall or reduced water demand (Newton, 1974; Owen, 1974; Wickens, 1983; Baum, 1996; Sorg & Rohner, 1996). The actual role of water storage in baobab trees, however, had not been examined until recently, and had simply been assumed from their physiognomy. In a related study we found no evidence that stored water was used
to maximize daily stomatal opening during the rainy season; instead, baobab trees appear to prioritize conduit safety and turgor maintenance (Chapotin et al., 2006). The goal of the present study was to understand the role of stem water during the dry season. Baobab trees, like all stem-succulent trees, are leaﬂess during most of the dry season, and new leaves are ﬂushed only towards the end of the dry season. As this phenological pattern considerably shortens their growing season, the ability to use stored water for early leaf ﬂushing could be advantageous. Stem water storage plays a physiological role in many different plants, and contributes to their ability to survive in a diverse range of ecosystems. Arborescent palms can use water stored in the stem to buffer xylem potentials and support leaves when water availability is limited (Holbrook & Sinclair,
1980.5 m from the ground. Borchert. 28 A. rubrostipa. Perrier. or simply by having the leaves present when the rainy season begins. Cedeño. and roots can access a large area around the tree. (Daubenmire. 1995). Borchert & Rivera. Baobabs range in height from 5 to 30 m.
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www. Access to the crown of the tree for installation of sap ﬂow probes and stomatal conductance measurements was through use of the single-rope technique (Laman.. Fewer studies have examined the role of water storage in deciduous trees (Nilsen et al. A second goal was to learn whether the use of stored water allows baobab trees to extend their growing season. Machado & Tyree..5 and 1. Lobo et al. 1996).. dried at 75°C and reweighed.. Phenology Vegetative phenology was assessed during several seasons between September 2001 and January 2003.
Madagascar and Australia (Armstrong. Borchert & Rivera.. and as baobab trees are thought to be shallow-rooted (Fenner. 1990. Their low-density wood has large vessels. Fenner. Soils are sandy with low water retention.. and abundant parenchyma throughout the stem (Fisher. while A. za are found over a large range of habitat types in western and southern Madagascar.550 Research
1992). 1995. Three cores were taken per site every 7–14 d from September to December 2003. Many deciduous. a pattern suggestive of a ‘droughtavoidance’ strategy (Owen. Waring et al. and Adansonia grandidieri Baill. while some tropical forest trees use stored water on a daily basis (Goldstein et al. Study trees ranged from 1 to 3.. with individual roots traveling along the surface for up to 70 m (Guy. 1994. 2001.5 m in diameter and 15–25 m in height. Contrary to expectation from their high water-storage capacity.5. Holbrook et al. 1980). 1995). 1994a. cylindrical or tapered trunks (Baum. 1990). any growth during this time should be dependent on stored water. 1996). 1971).. Newton. and the proportion of species with this growth form increases in drier areas (Medina. 1981. commonly referred to as baobabs. 1998).org © The Authors (2005). 1974. The soil cores were placed in sealed plastic bags in the ﬁeld.newphytologist. Additionally. which probably form a monophyletic group (Baum. Baobab trees. are restricted to the drier regions in the north. 1984). 1996). 2002). except near seasonal watercourses where increased clay content confers a higher water-retaining capacity (Sorg & Rohner. have a disjunct distribution across Africa. 1998). Three sections 5 cm in length and centered at 2. then brought to the laboratory. Our major objective in this study was to learn to what extent stored water is relied on during the dry season and for leaf ﬂushing. 1990). Stratton et al. Adansonia za Baill. ﬂush new leaves before the rainy season. and exhibit a wide variety in overall morphology. stem-succulent trees shed their leaves earlier than co-occurring. latitude 20°27′ S) (Rakotonirina. The majority of precipitation occurs during the rainy season (December–March) and rainfall is absent during the main part of the dry season (April–October). Study species The eight species in the genus Adansonia. 1980. The six Malagasy species. Leaf bud break and crown ﬂush during the 2001 and 2003 leaf-ﬂushing periods. transpiring conifers (Waring & Running. 1996. Baum. Journal compilation © New Phytologist (2005)
. as can large. west and south of the island. Rooting depth is generally shallow. za and nine A. 1995). Observations were made on 45 trees of A. 1995. weighed. often before or at the onset of the dry season. but can be highly variable and ranges from 300 to 1400 mm yr−1 (Ganzhorn et al. dry forest trees have large. Additional trees smaller than 1 m in diameter were included for phenological observations. were monitored on a weekly basis. Mean annual rainfall is 770 mm. Soil water content Soil cores were taken from a depth of 10–30 cm below the leaf-litter layer at four representative sites in the Kirindy Forest to quantify soil water content during the dry-to-rainyseason transition. Guy. swollen stems with a high water content (Holbrook et al. 1974). Stem water content Cores 12 mm in diameter and 50 cm in length were extracted from the stem. either by initiating stomatal opening before the onset of the rainy season. 2000.. swollen. stem succulents may close their stomata to prevent transpiration and turgor loss (Nilsen et al. Adansonia rubrostipa and A. where they are found in deciduous forests. 1995). like many stem succulents. 1971. 2001. Rivera et al. 2001). 1983). 1978. 15 and 30 cm beneath the bark were immediately
Materials and Methods
Field site This study was conducted at the Kirindy Forestry Reserve. nonsucculent trees. (the latter was included only for measures of water content and phenology). during short periods of drought... The end of the dry season is when soil water availability is lowest. between 0. & H. scrublands and savannas. grandidieri has a narrower range and a more consistent morphology (Baum. 1972. Fenner.. spiny forests. 2003). and extent of leaf drop during the 2002 leaf-shedding period. Water content was calculated as a percentage of dry weight. Holbrook.. 1979) and Andean rosette species (Goldstein et al. 1994b. few ﬁbers. Frankie et al. grandidieri. and all have huge. 1974. The three species of baobab examined in this study were Adansonia rubrostipa Jum. one of the largest remaining tracts of tropical dry forest in western Madagascar (longitude 44°49′ E. using an increment borer.
5–1. 2002b). (2002). Two more measurements were made in September and December 2003.
Phenology Leaf break was initiated in early October and leaf shedding in early March. Cores were extracted from ﬁve trees each of A. (1999) discuss the implications of using sap ﬂow probes longer or shorter than conductive sapwood depth and the various corrections that may be applied when the exact sapwood depth
Table 1 Dimensions of main stem and small branches at the location of probe insertion for each tree on which sap ﬂow measurements were made Parameter Diameter at base (cm) Diameter at top (cm) Diameter of branch 1 (cm) Diameter of branch 2 (cm) Height to main branches (m) AR1 AR2 AR3 AZ1 AZ2 AZ3 127 80 10 13 13 147 92 11 11 12 110 116 8 8 11 119 82 11 9 13 128 94 10 11 13 112 87 11 13 17
Adansonia rubrostipa (AR). Total water content in the stems of several representative trees at different times of year was estimated by taking the diameter of the tree at several points along the stem and the height at the point of branch divergence. Water content was calculated as percentage of fresh volume (Domec & Gartner. measurements on AR3 and AZ3 during the transitional period between leaf-ﬂushing period and rainy season. Probes were inserted to a depth of 1 cm beneath the bark. using 12–15 leaves per tree. Five additional trees of A. Adansonia za (AZ). February and March 2003. In the absence of accurate sapwood depth estimations. Permanent guides were used to ensure repeated measurements with the dbh tape were made at the same height each time. representative morning time-courses were made on newly ﬂushed leaves from one tree of each species. za trees was measured with a Li-Cor 1600 steady-state porometer. rubrostipa. Measured leaves were distributed. grandidieri four times between October 2001 and February 2002. za were included to increase the sample size for these last two measurements. Clearwater et al. za. Further measurements were taken in two separate years (2001 and 2003) during the dry season to compare levels of stomatal conductance before and after the rainy season began. The probes were therefore heated discontinuously. which led to signiﬁcant errors when the traditional Granier calibration was applied to continuously heated probes. throughout the outer crown of the tree. 2002).
is known. Journal compilation © New Phytologist (2005) www. and were placed at the base of the trunk (0. rubrostipa and one A. AZ1 and AZ2 were made during the leaf-ﬂushing period. Stomatal conductance Stomatal conductance of A. according to the formula: ASD = es − ea where es is the saturated water vapor pressure and ea is the ambient water vapor pressure of the air. 1985). and we present our data as sap ﬂux densities. we focus on the relative timing and overall pattern of sap ﬂow at the different points in the tree. Unheated reference probes revealed large underlying temperature gradients in the stem and branches. to the extent possible given the inherent difﬁculties with access. Stem diameter at breast height was measured monthly with a dbh tape to assess stem shrinkage and swelling in response to uptake or use of stem water. and calculated sap ﬂow rates from probes at the same location were averaged. AR2. rubrostipa and A. rubrostipa and A.Research
sealed into preweighed plastic bags. and the average value calculated.0 m above ground) to measure sap ﬂow from the roots to the stem. at the top of the trunk (within 1 m of the point of main branch divergence) to measure sap ﬂow from the main stem to the branches. Our measurements indicated that the conductive sapwood in some places was probably < 1 cm in depth. and rates of sap ﬂow were calculated by taking the difference between heated and unheated measurements and then applying their calibration formulas. Ten to 15 leaves per tree were measured at each point in time. Two probes were installed at each location. Sap ﬂow rates were measured in two trees of each species during each measurement time (leaf-ﬂushing period and transitional period). it was not possible to resolve the actual conductive sapwood depth. which we estimated by placing probes at different depths to measure presence or absence of sap ﬂow at that point. with probes similar in design to those described by James et al. These measurements were made between 09:00 and 10:00 h on three trees of each species. following Do & Rocheteau (2002a. Sap ﬂow measurements Rates of sap ﬂow were measured using the thermal dissipation technique (Granier.newphytologist. Measurements were taken every 30–60 min from dawn to dusk to quantify the diurnal pattern of stomatal conductance during the 2002 rainy season ( January and February) in three trees of A. then again in January. Fresh weight and volume were determined before drying the cores at 75°C to a constant weight (24 h). and using the above stem water content values. A. Air saturation deﬁcit (ASD) at each measurement time was calculated from the relative humidity and temperature of the air as measured by the porometer. Sap ﬂow measurements on AR1.org
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. During the 2003 dry season. As probe tips were themselves 1 cm in length. za and A. All trees had a full crown of mature leaves by
© The Authors (2005). and on distal branches (8–13 cm in diameter) to measure sap ﬂow from the branches to the leaves. Dimensions of the stem and branches at the locations of probe insertion are presented in Table 1.
sap ﬂow rates at the bottom of the stem were low. The probes at the top of the stem had the highest sap ﬂow rates of the three positions. Adansonia grandidieri.1 to 8.4 and 12. rubrostipa. and were completely leaﬂess by mid-April (Fig. n = 9). The total volume of water lost was calculated for representative A. and did not begin to increase until after signiﬁcant rainfall had occurred. shrank considerably during the leaf-ﬂushing period compared with the dry season (Fig.newphytologist.2% were measured during the leaf-ﬂushing period (September– December) in two separate years (Fig. Stem shrinkage in A. rubrostipa. rubrostipa. Sap ﬂow rates were highest in the morning. Positive sap ﬂow rates through the upper stem and branches while rates at the base of the tree were close to zero conﬁrmed that water reserves in the stem were used to support leaf ﬂushing (Fig. then gradually decreased after the leaves were expanded. a greater height. (b) Daily rainfall during the same periods. za. During the leafﬂushing period. 1 (a) Percentage of study trees of each species with a full crown of leaves in two separate seasons (Adansonia rubrostipa. top and branches) to increase. the total volume of water for similarly sized trees of each species was approximately the same (1000 l). when ASD of the air was lowest. 4). rubrostipa. rubrostipa and A. although sap ﬂow rates decreased quickly as soil water was depleted (Fig. After the onset of the rainy season. Note that the last measurements (28 December 2003) were made at the beginning of the rainy season and do not reﬂect the maximal water content values that are expected once the surface soil water has replenished fully. n = 28.4 at the sites measured (percentage of dry weight. Journal compilation © New Phytologist (2005)
. Adansonia za. grandidieri.org © The Authors (2005). n = 45. and to a lesser extent A. grandidieri. water content values ranged from 3. but decreases in stem water content of 10. grandidieri.8 ± 0.to ﬁvefold greater volume of water loss for each meter of stem height than in A. za trees. sap ﬂow rates at all points in the tree reached higher values and
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www. Measurements of total stem volume were not available for A. Despite the fact that the majority of water lost was through shrinkage for A. coincident with greater soil water availability. stem volumetric water content did not change signiﬁcantly during the dry season ( March–September).8 ± 0. There was a slight decrease in stem water content in A. 3). Occasional rainfall events after the leaves had expanded caused sap ﬂow rates at all points in the tree (base. mean ± SE). and less taper than A. but the difference was not signiﬁcant. but A. grandidieri would correspond on average to a four. All species experienced a very small decrease in diameter during the dry season. 5). but remained constantly low within each site from September to December. indicating little to no water entry from the roots and soil into the stem during this time. za
and through water extraction with little size change for A. 2). za and A. 1). Water-use rates in the branches and the top of the stem increased as leaf ﬂushing progressed. which generally has a larger diameter.
mid-December. and declined to lower values over the course of the day. Soil water content Soil water content varied between the different sites at the Kirindy Forest.552 Research
Fig. Sap ﬂow rates During the leaf-ﬂushing period. and those in the small branches had moderate rates. Stem water content In A.
Adansonia grandidieri (AG). 2 Stem water content at three depths in the tree before and after the dry season (left). 7). (b) Adansonia za (AZ). (a) Adansonia rubrostipa (AR). not all trees exhibited early morning stomatal opening. Values from before the dry season are maximum values in stem water content obtained at the end of the rainy season. although the degree of stomatal opening was lower than during the rainy season by over an order of magnitude.05). Adansonia za (AZ). Leaf ﬂushing in certain forests
Fig. but stomata closed as the morning progressed and air-saturation deﬁcit increased (Fig. 6).5–1 l dm−2 h−1 at base. Stomatal conductance Stomatal conductance during the leaf-ﬂushing period was close to zero and remained low until after the rainy season began (Fig. and in some cases values of stomatal conductance were zero even during the morning after leaves were fully ﬂushed (data not shown). Values are mean of ﬁve trees for all species during the dry season and AG during the leaf-ﬂushing period. Values are mean diameter decrease ± 1 SE. Chapotin.
Phenological patterns in African tropical dry forests vary considerably from site to site. P < 0. 2005). A paired t-test (P < 0.
generally remained high throughout the day (0. 3 Diameter decrease at breast height during the dry season and leaf-ﬂushing period for Adansonia rubrostipa (AR). and for 10 trees of AR and AZ during the leaf-ﬂushing period ± 1 SE. and before and after the leaf-ﬂushing period (right). However.05) indicated signiﬁcant differences in stem diameter before and after each time period for AR and AZ during the dry season.org
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.newphytologist. Journal compilation © New Phytologist (2005) www. (c) Adansonia grandidieri (AG). 2–3 l dm−2 h−1 at top.
© The Authors (2005).Research
Fig. Some trees exhibited a small amount of stomatal opening shortly after sunrise. An asterisk between two data points indicates a signiﬁcant difference over that time period (paired t-test. and for all species during the leaf-ﬂushing period.
data are not available for any top probe on the second AZ tree).org © The Authors (2005).newphytologist.
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Fig. 4 Sap ﬂow rates in (a) two Adansonia rubrostipa (AR) trees. Journal compilation © New Phytologist (2005)
. Each line is the average of values from two individual probes in similar locations (except one probe only for the top probe on the ﬁrst AZ tree. Leaves are fully ﬂushed by the times indicated by arrows. (b) two Adansonia za (AZ) trees during the leaf-ﬂushing period.
relying on stored water might allow baobab trees to maintain stomatal opening even when soil water is unavailable. Stem water usage Stem water content was high in all species (71–75% by volume in the outer sapwood).org
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. and we therefore hypothesized that the water required for early leaf ﬂushing and transpiration in baobab trees derives from stored reserves in the tree. Journal compilation © New Phytologist (2005) www. 5 Sap ﬂow rates in (a) one Adansonia rubrostipa (AR) tree. while in others. Increasing water use during leaf ﬂushing and decreasing water use thereafter probably reﬂect changing leaf area and cuticular properties of the leaves during maturation (Fig.Research
© The Authors (2005). 4). The end of the dry season is generally when soil water content is at its lowest and evaporative demand at its highest.5 to 8. Leaves were fully ﬂushed by the time measurements were initiated. Sap ﬂow rates responded quickly to small rainfall events (arrows) ranging from 1. water-content values in baobab trees (500– 800%) are much higher than published values for other tree species (30–300%. although it should be noted that these values are high also because the solid fraction in baobab stems is quite low (Chapotin. and not from the soil. and generally decreased with
increasing depth into the stem. 1982). thereby increasing the length of the growing season. such as the Kirindy Forest.
can be entirely opportunistic with respect to water availability (Lieberman.5 mm total precipitation. Rainfall amount (mm) given next to each arrow. The leaf-ﬂushing period in baobab trees at Kirindy occurs towards the end of the dry season. 1994a). Tsoumis. The measured sap ﬂow patterns were consistent with the hypothesis that the water necessary for leaf ﬂushing was derived almost entirely from the stem and branches. Furthermore. 1991) as well as other stem-succulent trees (120–410%. and a full crown of mature leaves is often maintained for several weeks before the onset of the rainy season. Each line is the average of values from two individual probes in similar locations (except one probe only for top of AZ).newphytologist. many trees ﬂush new leaves before the rainy season (Sorg & Rohner. Borchert. and moderate ﬂow rates in smaller branches reﬂect water moving into the leaves to support leaf expansion and cuticular water loss. 2005). When expressed as a percentage of dry weight. The presence of sap ﬂow at the top of the stem indicated substantial amounts of water moving from the stem into the branches. 1996). (b) one Adansonia za (AZ) tree during the transitional period between dry and rainy seasons.
just after the rainy season has begun (January 2004). as even closely related species can differ greatly in wood composition and the extent of shrinkage with water loss. and thus stomata must also be independently regulated by another factor (Fig. These results underscore the importance of not relying solely on water content or diameter changes when evaluating stem water loss. This was similar to the pattern observed in another dry forest deciduous tree. 6).
Adansonia rubrostipa and A. after leaves are fully mature (December 2003). We propose that limited water availability in the upper soil layers during this time restricts stomatal opening and. alternating with bands comprised primarily of vessels and ﬁbers. the grouping of the parenchyma cells into tangential bands. This increase in water ﬂow was noticeable at all points measured in the tree. za differed in the degree to which stem diameter shrank with water withdrawal. allowed the stem to shrink when water was withdrawn
Fig. rubrostipa). Journal compilation © New Phytologist (2005)
.05). Stem water does not support stomatal opening There was no evidence that stem water reserves were being used to support stomatal opening. Comparing the relationship between stomatal opening and ASD during the dry season with that during the rainy season demonstrates that greater stomatal closure is not simply a factor of increased ASD during the dry season. In A. with ﬁve to 15 leaves measured per tree.
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www. Air saturation deﬁcits (ASD) during each measurement period are also shown. although both species lost a comparable volume of water during this period. Difference between the values marked by a′ and b′ had only a tendency towards signiﬁcance (P = 0. P < 0. Values are mean of 10 –15 leaves per tree ± 1 SE. photosynthetic activity. 5). Signiﬁcant differences between months within a species are indicated by different letters (t-test.0 MPa during the afternoon. za in the rainy season of 2002). za. Values are mean of three trees per species ± 1 SE (except one tree only for A. however. 2002). and at the base of the stem. In A. vessels and parenchyma cells appeared to maintain the volume of the wood as air replaced withdrawn water. as stomatal conductance was consistently low until after the rainy season began and soil water was replenished (Fig. Measured leaf-water potentials before the rainy season reached a minimum of c.org © The Authors (2005). 7 Stomatal conductance during early morning on newly ﬂushed leaves for one Adansonia rubrostipa and one Adansonia za tree in December 2003. The rapidity with which water uptake followed individual rainfall events is signiﬁcant in that it indicates that ﬁne roots were present by the time leaves were ﬂushed or produced quickly upon soil
Fig. −1. and in the middle of the rainy season (January/ February 2002). where electron-transfer rates in newly ﬂushed leaves were low until after the rainy season began (Brodribb et al. as a consequence. All measurements were made between 09:00 and 10:00 h. Enterolobium cyclocarpum. which was near the turgor-loss point for leaves (unpublished data for A.newphytologist. 8). rubrostipa the uniformly distributed ﬁbers. 6 Leaf stomatal conductance in Adansonia rubrostipa and Adansonia za during leaf ﬂushing (November 2003).06). Whole-tree water use increased rapidly after isolated rainfall events (Fig.556 Research
without an increase in the air fraction. but particularly so in the small branches as a result of increases in transpirational water loss from the leaves. indicating the seasonal onset of water uptake by the roots..
Each point is the mean of ﬁve to 15 leaves from one tree ± 1 SE.org
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. 1988. By restricting water use to leaf ﬂushing rather than stomatal opening. daily temperatures reach their yearly maximum during this time. Nontranspiring canopy leaves frequently reached 40°C. and the inability to reduce leaf temperatures through evaporative cooling exposes the leaves to potentially damaging temperatures. Having the leaves present by the end of the dry season allowed baobab trees to open their stomata as soon as soil water became available.. the constraints described above. The timing of the rainy season can vary by several months. and the total amount of precipitation in any given rainy season is similarly variable (Fenner. Furthermore. 1980). indicating transpirational water loss caused by stomatal opening. Baobab stem wood is highly vulnerable to cavitation and has a turgor-loss point near actual ﬁeld water potentials (Chapotin et al. that sap ﬂow and stomatal conductance reached their maximal values and remained consistently high (Chapotin et al. Sap ﬂow rates then remained low until the next rainfall event. 8 Stomatal conductance as a function of air saturation deﬁcit (ASD) in Adansonia rubrostipa trees during the dry and rainy seasons. Journal compilation © New Phytologist (2005) www. may require that use of stored water be limited in this way. In an environment where the rainy season is short and its length can vary by several months. With regard to the second question.. This raises two main questions: why does leaf ﬂushing in baobab trees precede the onset of the rainy season. but also in the branches. when rainfall events were frequent enough to prevent soil drying. given the vast amounts of water in the trunk. although they may be of limited duration or intensity. This rate of water movement may be sufﬁcient for the needs of growing tissues. not only at the base of the tree. the baobab tree is able to maintain leaf turgor until the onset of the rainy season. however. 2006). a temperature that induces heat-stress responses in many plants ( Jones. the wood is soft and weak. early leaf ﬂushing allows baobab trees to take advantage of scattered or early rainfall events. as to why more extensive use of stored water does not occur.newphytologist. 2005). but is unlikely to meet the demand of actively transpiring leaves. Three to ﬁve trees per curve. and why is stored water not drawn upon more extensively to maximize carbon assimilation? The ﬁrst question is pertinent because the use of stored water for leaf ﬂushing appears to necessitate some investment in water-storage tissue.
hydration. 2006). 1992). as they were in another dry forest site (Kummerow et al. and to produce a new crop of leaves the following year if current-year precipitation levels were not
© The Authors (2005). and the highly unpredictable climate to which baobab trees are subject. results from two related studies indicate that stored water may be unavailable because of tissue water relations. 1990). Limited use of stem water despite early ﬂushing Early leaf ﬂushing in baobab trees does not appear to allow stomata to open before the rainy season when soil water availability is still low. It has been suggested that a physiological advantage of early leaf ﬂushing is that leaves are ready to begin photosynthesizing when the rainy season begins (Rivera et al. and comprised primarily of living tissues which may exhibit decreased mechanical stability under excessive water withdrawal (Niklas. However. It was not until after the onset of the rainy season.. These increased rates of water use were not sustained. transport limitations and biomechanical considerations.. maximizing photosynthesis in this way would be advantageous. indicating water uptake from the soil. in response to declining soil water availability. In addition. and rates of sap ﬂow in the branches and stem declined gradually over the several days following the rainfall event. All measurements were made between 07:00 and 15:30 h. and is likely to occur very slowly. 2002). Chapotin. the conductive portion of the sapwood is restricted to the 1–2 cm just beneath the bark.Research
Even very small rainfall events occurring before the full onset of the rainy season were accompanied by immediate increases in sap ﬂow. Furthermore. so that radial transport of water in the stem to the transpiration stream is through a relatively high resistance pathway. Additionally. Water use in baobab trees before the onset of the rainy season appears surprisingly conservative. and eliminated the time lag that would otherwise have existed between the onset of the rainy season and the onset of photosynthetic activity.
Chapotin S. ecology and habitat. Meinzer FC. Meinzer FC. Primate Report 44: 311–328. Systematic Biology 47: 181–207.558 Research
sufﬁcient to replenish the water content in the wood. Almacenamiento de recursos y relaciones hídricas durante el desarrollo ontogenético de Cochlospermum vitifolium. Wound healing by exposed secondary xylem in Adansonia (Bombacaceae). Maas M. 1974. Licenciada Thesis. 2. Hydraulic and photosynthetic co-ordination in seasonally dry tropical forest trees. 1971. Do F. Vegetatio 90: 73–80. Jackson P. 2nd edn. Plant. Quesada M. Bombacaceae) as inferred from multiple data sets. Goldstein G. 2002a. Cell & Environment 15: 401–409. Rocheteau A. Cavelier J. IAWA Bulletin 2: 193–199. 1972. Holbrook NM. Some measurements on the water relations of baobab trees. In: Gartner BL. Lieberman D. Tree Physiology 22: 641–648. Annales des Sciences Forestières 42: 193–200. 1994b. Gutiérrez MV. How do water transport and water storage differ in coniferous earlywood and latewood? Journal of Experimental Botany 53: 2369–2379. Gartner BL. The results of this study appear to contradict general ideas about water use in baobab trees by indicating a limited role for stored water. Potential errors in measurement of nonuniform sap ﬂow using heat dissipation probes. (Bombacaceae). Clearwater MJ. Sinclair TR. Rivera G. We also thank three reviewers for their helpful comments and suggestions. Herrerías-Diego Y. Safety recommendations for climbing rain forest trees with ‘Single Rope Technique’. CA. Inﬂuence of natural temperature gradients on measurements of xylem sap ﬂow with thermal dissipation probes. Journal of the Botanical Society of South Africa 57: 30–37. una especie pionera del Bosque Seco de Costa Rica.
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This project was made possible through funding by the Sinclair Kennedy Traveling Fellowship. The ecology and conservation of the baobabs of Madagascar. Whitbeck JL. Oecologia 84: 126–133. II. Cambridge. Tree Physiology 22: 649–654. the results suggest that water-use patterns in stem-succulent trees may need further study. Larigauderie A. 1981. 1994a. 1982.x Clearwater MJ. 1992. 1995. Borchert R. The disjunct distribution of the genus Adansonia L. Cell & Environment 7: 179–186. Tree Physiology 22: 277–283.org © The Authors (2005). Fuchs EJ. Granier A. Heather Patt and Mario Ramohavelo provided ﬁeld assistance. Plant. Borchert R. Stoner KE. Goldstein G.01456. Celis A. 1995. Comparative phenological studies of trees in tropical wet and dry forests in the lowlands of Costa Rica.newphytologist. Abraham J-P. Ecology 75: 1437–1449. Rodolphe Rasoloarivony. Stem-succulent trees occur in many distantly related plant taxa. (In press. Biogeography and ﬂoral evolution of baobabs (Adansonia. Biotropica 12: 205–209. Biotropica 27: 406–409. Daubenmire R. 2002. Journal of Ecology 60: 147–170. 1983. Rojas J. Annals of the Missouri Botanical Garden 82: 440 – 470. Transpiration and stem water storage. Holbrook NM.1111/j. 151–174. 1998. 2005. To our knowledge there are no similar studies on stem-succulent trees that focus speciﬁcally on water use during the period between leaf ﬂushing and the onset of the rainy season. Borchert R. 1980. 1990. Journal of Ecology 70: 791–806. Plant. Ganzhorn JU. 1985. USA: Academic Press. Kummerow J. Field observations and possible remedies. Tropical Dry Forests. Universidad de Costa Rica. Goldstein G.2005. Plant Stems: Physiology and Functional Morphology. Holbrook NM. Ramananjatovo A. Chapotin S. Trees 8: 115 –125. Soil and stem water storage determine phenology and distribution of tropical dry forest trees. Mooney HA. Stem water storage and diurnal patterns of water use in tropical forest canopy trees. Although this study focuses on just one genus. Lobo JA. Holbrook NM. Mooney HA. Inﬂuence of natural temperature gradients on measurements of xylem sap ﬂow with thermal dissipation probes. Holbrook NM. PhD Thesis. Seasonality and phenology in a dry tropical forest in Ghana. 1990. Cell & Environment. Holbrook NM. James SA.) doi:10. Opler PA. Harvard University. 1. the Arnold Arboretum and the Department of Organismic and Evolutionary Biology at Harvard University. 1995. Phenology and other characteristics of tropical semi-deciduous forest in north-western Costa Rica. Jones HG. Andriamanarivo L. Drought responses of neotropical dry forest trees. Fisher JB. 1996. Medina E. Ganzhorn AW. Baker HG. Cell & Environment 25: 1435 –1444. Water balance in the arborescent palm. 2002. Cell & Environment 21: 397–406. Water status and development of tropical trees during seasonal drought. MA. Cambridge. Frankie GW. The baobabs: Adansonia spp. ed. Brodribb TJ. Andrade JL. Journal compilation © New Phytologist (2005)
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Climate and tree phenology of the dry deciduous forest of the Kirindy Forest. 1996. Journal compilation © New Phytologist (2005) www. Is the baobab tree succulent? Cactus and Succulents Journal 36: 57–58. Shariﬁ MR. A contribution to the ecology of the African baobab (Adansonia digitata L. Primate Report 46: 57–80.uk. The baobab – Africa’s upside-down tree. Whitehead D. In: Bullock SH. Rapid reports and both Modelling/Theory and Methods papers are encouraged. Complete information is available at www. Owen J. Rakotonirina. Increasing day-length induces spring ﬂushing of tropical dry forest trees in the absence of rain. eds. go to the website and click on ‘Journal online’. 1995. Meinzer FC.
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Bombacaceous trees in seasonal forests in Costa Rica and Mexico. Cell & Environment 1: 131–140. Dependency of the tensile modulus on transverse dimensions. and we provide 25 offprints as well as a PDF for each article.gov. The contribution of stored water to transpiration in Scots pine. Stratton L. Plant. USA: Chapman & Hall. Oecologia 82: 299–303. Waring RH. American Journal of Botany 75: 1286 –1292. 2002. Newton L. do get in touch with Central Ofﬁce (newphytol@lancaster. Nilsen ET. Seasonally Dry Tropical Forests. Ehleringer JR. Water relations of stem succulent trees in north-central Baja California. 221–242. 1983. Plant.). Trees 16: 445–456. water potential. Sapwood water storage: its contribution to transpiration and effect upon water conductance through the stems of old-growth Douglas-ﬁr. 1990. Rivera G et al. • Regular papers.org
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© The Authors (2005). Letters. • For online summaries and ToC alerts. Jarvis PG. 1974. You can take out a personal subscription to the journal for a fraction of the institutional price. Machado J-L. Rohner U. and cell number of pith parenchyma. Goldstein G. the US Ofﬁce (newphytol@ornl. from online submission through to publication ‘as-ready’ via OnlineEarly – the 2004 average submission to decision time was just 30 days. Mooney HA. 2000. Research reviews.newphytologist. 1988. Forseth IN. Composition and structure of a dry forest on sandy soils near Morondava. 1979. 1996. Wickens GE. for a local contact in North America. Savanna 3: 1–12. 1994. Stem water storage capacity and efﬁciency of water transport: their functional signiﬁcance in a Hawaiian dry forest. Science and Technology of Wood. American Journal of Botany 90: 1054 –1063. Primate Report 46: 81–88. Online-only colour is free. 1991. Cell & Environment 2: 309–317.