Changes in gas exchange characteristics and water use efficiency of mangroves in response to salinity and vapour pressure deficit

Summary Measurements were made of the photosynthetic gas exchange properties and water use efficiency of 19 species of mangrove in 9 estuaries with different salinity and climatic regimes in north eastern Australia and Papua New Guinea. Stomatal conductance and CO₂ assimilation rates differed significantly between species at the same locality, with the salt-secreting species, Avicennia marina, consistently having the highest CO₂ assimilation rates and stomatal conductances. Proportional changes in stomatal conductance and CO₂ assimilation rate resulted in constant and similar intercellular CO₂ concentrations for leaves exposed to photon flux densities above 800 mol·m^-2·s^-1 in all species at a particular locality. In consequence, all species at the same locality had similar water use efficiencies. There were, however, significant differences in gas exchange properties between different localities. Stomatal conductance and CO₂ assimilation rate both decreased with increasing salinity and with increasing leaf to air vapour pressure deficit (VPD). Furthermore, the slope of the relationship between assimilation rate and stomatal conductance increased, while intercellular CO₂ concentration decreased, with increasing salinity and with decreasing ambient relative humidity. It is concluded from these results that the water use efficiency of mangroves increases with increasing environmental stress, in this case aridity, thereby maximising photosynthetic carbon fixation while minimising water loss.Contribution No. 459 from the Australian Institute of Marine Science     

Photosynthesis of mistletoes in relation to their hosts at various sites in tropical Brazil

 Chlorophyll a fluorescence parameters showing the instantaneous performance and carbon-isotope ratios reflecting long-term behaviour of leaves were determined for a large number of mistletoe/host-pairs in the cerrado belt of Brazil. Study sites were a very exposed rupestrian field, a semi-exposed savanna and a highly shaded gallery forest. The major question asked was if photosynthetic capacity of mistletoe leaves differed from that of the leaves of their respective hosts. It is shown that except for the very exposed rupestrian field site, photosynthetic capacity appeared to be similar in mistletoes and host leaves. The superior behaviour of host leaves in the rupestrian field was due to particularly expressed sun-plant characteristics of the host. However, mistletoes always had higher average stomatal conductances, lower leaf temperatures at similar or even higher irradiance and higher intercellular CO₂-partial pressures than hosts. Photosynthetic performance of mistletoe leaves was independent of whether a given mistletoe species parasitized aluminium-accumulating or non-accumulating host species in the cerrados with their aluminium-rich soils. Received: 7 April 1997 / Accepted: 20 August 1997     

Comparative ecophysiology of five species of Sedum (Crassulaceae) under well-watered and drought-stressed conditions

Summary Gas exchange patterns, diurnal malic acid fluctuations, and stable carbon isotope ratios of five species of Sedum were investigated to assess the ecophysiological characteristics of three different photosynthetic pathways under well-watered and drought-stressed conditions. All five species have succulent leaves and stems and were examined under identical environmental conditions. When well-watered, Sedum integrifolium (Raf.) Nels. and S. ternatum Michx. displayed C₃ photosynthesis, S. telephioides Michx. and S. nuttallianum Raf. exhibited CAM-cycling, and S. wrightii A. Gray showed CAM. When grown under a less frequent watering regime, S. integrifolium and S. ternatum exhibited CAM-cycling, whereas S. telephioides and S. nuttallianum displayed CAM-cycling simultaneously with low-level CAM. Sedum wrightii retained its CAM mode of photosynthesis. In general, leaf ¹³C values reflected these variations in photosynthetic pathways. While all values of water-use efficiency (WUE) were greater than those reported for most C₃ and C₄ species, no correlation of malic acid accumulation in the CAM and CAM-cycling (including low-level CAM) species with increased WUE was found. Sedum wrightii (CAM) had the highest WUE value at night, yet its 24-h WUE was not different from S. ternatum when the latter was in the C₃ mode. Thus, relative water-use efficiencies of these species of Sedum were not predictable based on photosynthetic pathways alone.     

Pioneer and late stage tropical rainforest tree species (French Guiana) growing under common conditions differ in leaf gas exchange regulation,carbon isotope discrimination and leaf water potential

Leaf gas exchange rates, predawn _wp and daily minimum _wm leaf water potentials were measured during a wet-to-dry season transition in pioneer (Jacaranda copaia, Goupia glabra andCarapa guianensis) and late stage rainforest tree species (Dicorynia guianensis andEperua falcata) growing in common conditions in artificial stands in French Guiana. Carbon isotope discrimination () was assessed by measuring the stable carbon isotope composition of the cellulose fraction of wood cores. The values were 2.7 higher in the pioneer species than in the late stage species. The calculated time integratedC _i values derived from the values averaged 281 mol mol^–1 in the pioneers and 240 mol mol^–1 in the late stage species. The corresponding time-integrated values of intrinsinc water-use efficiency [ratio CO₂ assimilation rate (A)/leaf conductance (g)] ranged from 37 to 47 mmol mol^–1 in the pioneers and the values were 64 and 74 mmol mol^–1 for the two late stage species. The high values were associated—at least inJ. copaia—with high maximumg values and with high plant intrinsinc specific hydraulic conductance [Cg/(_wm–_wp], which could reflect a high competitive ability for water and nutrient uptake in the absence of soil drought in the pioneers. A further clear discriminating trait of the pioneer species was the very sensitive stomatal response to drought in the soil, which might be associated with a high vulnerability to cavitation in these species. From a methodological point of view, the results show the relevance of for distinguishing ecophysiological functional types among rainforest trees.     

Stem and leaf water potentials, gas exchange, sap flow, and trunk diameter fluctuations for detecting water stress in lemon trees

The sensitivity of continuous (on a whole-day basis) and discretely (at midday) measured indicators of the plant water status in adult lemon trees in response to a cycle of water deprivation and recovery, and the feasibility of obtaining baselines for tree water status indicators was investigated in 30-year-old Fino lemon trees (Citrus limon (L.) Burm. fil.) grafted on sour orange (C. aurantium L.) rootstocks. Control plants (T0) were irrigated daily above their crop water requirements in order to obtain non-limiting soil water conditions, while T1 plants were subjected to water stress by withholding irrigation for 50 days, after which time irrigation was restored and plant recovery was studied for 16 days. In T0 plants the water relations and the plant symptoms confirmed that they had not suffered waterlogging. In contrast, T1 plants showed a substantial degree of water stress, which developed very slowly. Maximum daily trunk shrinkage (MDS) increased in response to water stress during the first 15 days of the experiment, but when the stem water potential (Ψ_stem) fell below −1.8 MPa, the MDS signal intensity decreased. However, Ψ_stem and sap flow (SF) signal intensities progressively increased during the water stress period. The results showed that MDS is a very suitable plant-based indicator for precise irrigation scheduling in adult lemon trees. Reference or baseline relationships for MDS, Ψ_stem, and SF measurements as a function of several parameters related to the evaporative demand of the atmosphere were obtained. This fact open up the possibility of considering a plant-based indicator measurement at a given time relative to the expected value under non-limiting water conditions, which can be calculated from the reference relationships.     

Comparative study of water uptake and photosynthetic gas exchange between scrub and fringe red mangroves,Rhizophora mangle L.

Summary The red mangrove (Rhizophora mangle L.) occurs frequently in both scrub and fringe mangrove forests. Our previous study demonstrated that individuals of this mangrove species growing in scrub and fringe forests differ significantly in both morphological and physiological characteristics. To further characterize physiological differences between scrub and fringe mangroves, we compared their differences in water uptake and photosynthetic gas exchange during different seasons. In the wet season (June–October, 1990), scrub mangroves showed lower D and ¹⁸O values of stem water than fringe mangroves, indicating more usage of rain-derived freshwater. In the dry season (Jan–April, 1991), however, scrub mangroves utilized the same water source as fringe mangroves, reflected by their similar D and ¹⁸O values of stem water. Consistently, there were significant differences in predawn water potentials between scrub and fringe mangroves in the wet season (October 1990) with higher values for scrub mangroves, but no significant differences in the dry season (January 1991). Higher elevation in the scrub forest seems to be the major factor responsible for the shift of water sources in scrub mangroves. On Apr. 27 and Aug. 8, 1990, scrub mangroves showed lower CO₂ assimilation rate, stomatal conductance, and intercellular CO₂ concentration than fringe mangroves. There were no differences in these gas exchange characteristics on the other two measuring dates: Oct. 17, 1990 and Jan. 11, 1991. Instantaneous water use efficiency was significantly higher for scrub mangroves than for fringe mangroves on three of the four sampling dates. Similarly, leaf carbon isotope discrimination of scrub mangroves was always significantly lower than that of fringe mangroves, indicating higher long-term water use efficiency. Higher water use efficiency in scrub mangroves is a result of stomatal limitation on photosynthesis, which may entail considerable carbon cost to the plants.     

Comparative gas exchange performance during the wet season of three Brazilian Styrax species under habitat conditions of cerrado vegetation types differing in soil water availability and crown density

We studied the influence of environmental conditions in different vegetation types of the Brazilian savanna (cerrado s.l.) on CO₂ assimilation and water use efficiency of Styrax ferrugineus, S. camporum and S. pohlii, which are morphologically similar but have distinct distribution patterns. We measured leaf gas exchange and water relations in adult and young plants. Adult plants of S. ferrugineus were measured in the cerrado s. str.; plants of S. camporum at the edge of cerradão, a woodland vegetation; and in a riparian forest those of S. pohlii. Eight-month-old young plants were planted in the cerrado s. str., at the edge and in the understory of the cerradão, and in the understory of the riparian forest. For young plants, the high light availability in the cerrado s. str. resulted in a threefold greater CO₂ assimilation rate (A) compared to the other sites. A of adult plants under full irradiation (1800 μmol photons m⁻² s⁻¹) was significantly lower in S. pohlii compared to S. ferrugineus and S. camporum. Although sufficient soil water was available at every site at the end of the wet season, the extreme high water content in the riparian forest soil led to increased stomatal conductance (gs) and transpiration rate (E) for young plants of S. camporum and S. pohlii, but not of S. ferrugineus. However, the large gs in S. pohlii and S. camporum young plants did not result in higher A rates. The differences may indicate some specialization of S. ferrugineus to best growth in open savanna and of S. pohlii in riparian forest habitats. The high water use efficiency of S. camporum young plants in the understory of cerradão seems not yet sufficient to emphasize that this species benefits from this particular habitat. Because, while water supply during the wet season is quite sufficient, soil water deficits and frequent fires can occur during the dry season and might modify strongly the habitat conditions for the three Styrax species.     

Photosynthetic gas exchange and the stable isotope composition of leaf water: comparison of a xylem-tapping mistletoe and its host

Photosynthetic gas exchange and the stable isotopic composition of foliage water were measured for a xylem tapping mistletoe, Phoradendron juniperinum, and its host tree, Juniperus osteosperma, growing in southern Utah. The observed isotopic composition of water extracted from foliage was compared to predictions of the Craig-Gordon model of isotopic enrichment at evaporative sites within leaves. Assimilation rates of juniper were higher and stomatal conductance was lower than the values observed for the mistletoe. This resulted in lower intercellular/ ambient CO₂ values in the juniper tree relative to its mistletoe parasite. For mistletoe, the observed foliage water hydrogen and oxygen isotopic enrichment was less than that predicted by the model. In juniper, foliage water hydrogen isotopic enrichment was also lower than that predicted by the evaporative enrichment model. In contrast, the oxygen isotopic enrichment in juniper foliage water was slightly greater than that predicted for the evaporative sites within leaves. Hydrogen isotopic enrichment in mistletoe foliage shows systematic variation with stem segment, being highest near the tips of the youngest stems and decreasing toward the base of the mistletoe, where isotopic composition is close to that of stem water in the host tree. In a correlated pattern, mid-day stomatal conductance declined abruptly in mistletoe foliage of increasing age.     

Gas exchange responses of a mangrove species, Avicennia marina, to waterlogged and drained conditions

This study was undertaken in summer on fully expanded leaves of Avicennia marina trees in the Beachwood Mangroves Nature Reserve, Durban,South Africa. Data sets were obtained over 5–7 days of relatively dry conditions and over two periods of 5 days during which the swamp was continuously inundated with dilute seawater (< 150 mol m⁻³NaCl). Gas exchange responses were strongly influenced by photosynthetic photon flux density (PPFD), leaf temperature and leaf to air vapour pressure deficit (Δw). Carbon dioxide exchange was saturated at a PPFD of about 800 μmol m⁻² s⁻¹. Maximal CO₂ exchange rates ranged from 8.5 to 9.9 μmol m⁻² s⁻¹ with no differences between drained and waterlogged conditions. Under drained conditions, leaf conductance,transpiration and internal CO₂ concentrations were generally lower, and water use efficiencies higher, than during waterlogging. Continuous waterlogging for 5 days had no adverse effect on CO₂ exchange. Xylem water potentials ranged from −1.32to −3.53 MPa during drained and from −1.02 to −2.65 Mpa during waterlogged conditions. These results are discussed in relation to anatomical and metabolic adaptations of A. marina to waterlogging stress. This revised version was published online in August 2006 with corrections to the Cover Date.     

Morphological and physiological variation in western redcedar (<Emphasis Type="Italic">Thuja plicata</Emphasis>) populations under contrasting soil water conditions

Adaptation to precipitation conditions may induce genetic diversity that changes morphological and physiological traits. This hypothesis was investigated in the seedlings of seven western redcedar (Thuja plicata Donn ex D. Don) populations, which were collected along a precipitation transect from the Pacific coast to the southern interior of British Columbia, Canada. The experimental seedlings were either well-watered or soil-droughted and measured for growth, gas exchange rates, transpiration efficiency, and carbon isotope discrimination during or at the end of the third growing season. Significant variation was found in most of these morphological and physiological traits among the populations. Much of this variation occurred under well-watered, but not so much under droughted conditions. Mean height increments and transpiration efficiency showed a significant linear relationship, but biomass increments exhibited a quadratic relationship with precipitation on the origin site of these populations. Measurements of water use efficiency obtained from instantaneous gas exchange measurements, carbon isotope discrimination, and transpiration efficiency were intercorrelated in the seedlings. However, neither did any of these measurements consistently rank the populations, nor were they indicative of adaptation to climatic precipitation conditions in these western redcedar populations.     

Seedling morphology of three mangrove species and its taxonomic implications

Seedling establishment is an important part of the life cycle of mangroves. Very early in the juvenile stage, they face adverse situations such as a variable and unstable substrate as well as a high-saline environment Therefore, the propagules should have some ability to float in saline water. Seedlings differ morphologically from the adult plants. Here, I examined the seedling growth stages in three species of mangrove:Aegialitis rotundifolia Roxb.,Aegiceras corniculatum (L) Blanco, andAvicennia marina (Forsk.) Vierh. that grow in the Sundarbans swamps of West Bengal. In all cases, fruits were the propagating units; seeds were inseparable from those fruits. Germination was rapid and required no dormancy period. The hypocotyi pierced the seed coat rather than emerging from the pericarp — a variation on the traditional viviparous type of germination. Seedlings then became exposed prior to abscission from parent plants. Based on germination patterns and differences in seedling morphology observed here, these species should be classified in a more relevant taxonomic group than was done previously.     

Shoot water relations and gas exchange of western hemlock and western red cedar seedlings during establishment on a reforestation site

Summary Shoot water relations, summer gas exchange response and morphological development of western hemlock [Tsuga heterophylla (Raf.) Sarg.] and western red cedar (Thuja plicata Donn) seedlings were monitored over the first growing season on a coastal reforestation site in British Columbia. In March, osmotic potential (_s) at saturation [_s(sat)] was –1.98 MPa and turgor loss point [_s(tlp)] –2.38 MPa for western hemlock, while western red cedar had –1.45 MPa _s(sat) and –1.93 MPa _s(tlp). Seasonally _s increased through June and then decreased through September, with western hemlock –0.15 to –0.50 MPa lower than western red cedar. Maximum bulk modulus of elasticity (_max) for western hemlock was 29.3 MPa in March, decreased to 15.0 MPa in June and increased to 25.0 MPa from July through September, while western red cedar _max was 10.6 MPa in March and around 8.0 MPa thereafter. Utilized turgor (T _util) for western hemlock was <40% from March through May, 69 to 78% from June through August and 96% in September, while western red cedar T _util was 68 to 73% during March and April, 84 to 96% from May through August and 100% in September. Maximum CO₂ assimilation rate (A) of western red cedar was more than double western hemlock, and for both species A declined in a linear fashion with increasing vapour pressure deficit (D). Maximum foliage conductance (g _wv) declined in a concave manner as D increased in both species, with western red cedar values 50 to 67% greater than western hemlock. Maximum daily g _wv declined in a concave manner as predawn shoot water potential (_pd) decreased, with maximum daily g _wv 1.8 to 3.6 times greater in western red cedar than western hemlock, when _pd was –0.25 and –1.4 MPa, respectively. Western red cedar, compared to western hemlock, had a greater increase in A as g _wv increased. Eight months after planting, western red cedar seedlings had twice the root growth, measured as root dry weight and root number, of western hemlock.     

Tissue water relations in elfin cloud forest tree species of Serrania de Macuira,Guajira, Colombia

Summary Diurnal courses of stomatal conductance, leaf water potential, and the components of tissue water potential were measured in six canopy species in an elfin cloud forest. High values of stomatal conductance were measured on cloudy days and during early morning and late afternoon of sunny days. Decreases in stomatal conductance with increases in vapour pressure deficit may have been a response to avoid further water deficits and suggested a stomatal response to changes in relative humidity. Daily transpiration varied between 470 and 1014 g m^-2 day^-1 during cloudy days and between 532 and 944 g m^-2 day^-1 during clear days. Stomatal conductance may have also responded to changes in leaf water potential, which was minimum at noon. The minimum tissue water potential measured in the field was -1.8 MPa in Myrcianthes fragrans, and the minimum turgor pressure was 0.49 MPa also in M. fragrans. There was a correlation between the osmotic potential and the minimum tissue water potential, suggesting that osmotic potential plays a major role in the maintenance of turgor in these species, in spite of the great variability in the elastic properties of leaf tissues. Turgor pressure decreased during the day following the course of water potential but never approached the turgor loss point, as it has been measured in some lowland rain forest trees. This is a strong indication that elfin cloud forest trees do not suffer severe water deficits, and that small tree stature is not directly related to water shortage.     

Extensive summer water pulses do not necessarily lead to canopy growth of Great Basin and northern Mojave Desert shrubs

Plant species and functionally related species groups from arid and semi-arid habitats vary in their capacity to take up summer precipitation, acquire nitrogen quickly after summer precipitation, and subsequently respond with ecophysiological changes (e.g. water and nitrogen relations, gas exchange). For species that respond ecophysiologically, the use of summer precipitation is generally assumed to affect long-term plant growth and thus alter competitive interactions that structure plant communities and determine potential responses to climate change. We assessed ecophysiological and growth responses to large short-term irrigation pulses over one to three growing seasons for several widespread Great Basin and northern Mojave Desert shrub species: Chrysothamnus nauseosus, Sarcobatus vermiculatus, Atriplex confertifolia, and A. parryi. We compared control and watered plants in nine case studies that encompassed adults of all four species, juveniles for three of the species, and two sites for two of the species. In every comparison, plants used summer water pulses to improve plant water status or increase rates of functioning as indicated by other ecophysiological characters. Species and life history stage responses of ecophysiological parameters (leaf N, ¹⁵N, ¹³C, gas exchange, sap flow) were consistent with several previous short-term studies. However, use of summer water pulses did not affect canopy growth in eight out of nine comparisons, despite the range of species, growth stages, and site conditions. Summer water pulses affected canopy growth only for C. nauseosus adults. The general lack of growth effects for these species might be due to close proximity of groundwater at these sites, co-limitation by nutrients, or inability to respond due to phenological canalization. An understanding of the connections between short-term ecophysiological responses and growth, for different habitats and species, is critical for determining the significance of summer precipitation for desert community dynamics.     

Living in a shallow burrow under a rock: Gas exchange and water loss in an Australian scorpion

The scorpion, Urodacus manicatus (Scorpionida: Urodacidae), inhabits temperate sclerophyll woodland in south-eastern Australia and excavates a shallow burrow into soil beneath a rock. Mean minimum and maximum temperatures within a burrow were respectively higher and lower than outside measurements, and relative humidity within a burrow remained higher than outside the burrow at all times. Using flow-through respirometry, VCO₂ and water loss were measured at three ecologically relevant temperatures (10, 20 and 30 °C) to assess patterns of gas exchange and the temperature dependence of VCO₂ and water loss rates. Daytime VCO₂ corresponds to a lower metabolic rate than reported for the closely related, but more arid region inhabiting species Urodacus armatus and Urodacus yaschenkoi. CO₂ output is continuous at 10 °C; however, at 20 °C scattered sharp depressions are observed and these become more frequent and periodic at 30 °C. Total water loss in U. manicatus is nearly double that in U. armatus, indicative of lessened selective pressure for water conservation within a mesic rather than xeric environment.     

Leaf functional traits of Neotropical savanna trees in relation to seasonal water deficit

The seasonal savannas (cerrados) of Central Brazil are characterized by a large diversity of evergreen and deciduous trees, which do not show a clear differentiation in terms of active rooting depth. Irrespective of the depth of the root system, expansion of new foliage in deciduous species occurs at the end of the dry season. In this study, we examined a suite of leaf traits related to C assimilation, water and nutrients (N, P) in five deciduous and six evergreen trees that were among the dominant families of cerrado vegetation. Maximum CO₂ assimilation on a mass basis (A_mass) was significantly correlated with leaf N and P, and specific leaf area (SLA; leaf area per unit of leaf mass). The highest leaf concentrations of both nutrients were measured in the newly mature leaves of deciduous species at the end of the dry period. The differences in terms of leaf N and P between evergreen and deciduous species decreased during the wet season. Deciduous species also invested less in the production of non-photosynthetic leaf tissues and produced leaves with higher SLA and maintained higher water use efficiency. Thus, deciduous species compensated for their shorter leaf payback period by maintaining higher potential payback capacity (higher values of A_mass) and lower leaf construction costs (higher SLA). Their short leafless period and the capacity to flush by the end of the dry season may also contribute to offset the longer payback period of evergreen species, although it may involve the higher cost of maintaining a deep-root system or a tight control of plant water balance in the shallow-rooted ones.     

Carbon and water balance in Polylepis sericea,a tropical treeline species

Polylepis sericea trees grow well above the continuous forest line in the Venezuelan Andes. In these environments, extreme daily temperature ranges can occur at any time of the year and trees experience a 4 month dry period. The purpose of this work was to study carbon and water relations of this species in the field during wet and dry seasons in order to understand this species' success at such high altitudes. Leaf gas exchange (portable system in open mode) and leaf water potential (pressure chamber) were measured at 1–2 h intervals during several daily courses at 4000 m elevation in the Páramo de Piedras Blancas. CO₂ assimilation versus leaf temperature curves were also obtained for this species in the laboratory. Clear differences in the measured parameters were observed between seasons. For a wet season day, maximum CO₂ assimilation rate was 7.4 mol m^-2 s^-1 and leaf conductance was relatively constant (approximately 100 mmol m^-2 s^-1)In the dry season day, maximum CO₂ assimilation rate was 5.8 molm^-2 s^-1 and leaf conductance was close to 60 mmolm^-2 s^-1. Minimum leaf water potentials measured were -1.3 MPa for the wet and -2.2 MPa for the dry season. The CO₂ assimilation-leaf temperature relationship showed a 13.4°C leaf temperature optimum for photosynthesis with maximum and minimum compensation points of 29.5 and -2.8°C, respectively. Maximum night-time respiration was relatively high (2.7 (imol) m^-2 s^-1)Our results show thatP. sericea maintains a highly positive carbon balance through all daily courses, even though there is a slight water stress effect during the dry season; this suggests that its carbon assimilation machinery is well adapted to the low temperatures and seasonal water stress found in the high tropical mountains.     

Relation of water transport to leaf gas exchange properties in three mangrove species

Mangrove species more tolerant to salinity may function with less efficient water transport, which may be related to more conservative water use. To test the hypothesis, we investigate the gas exchange and hydraulic properties of three mangrove species: Rhizophora mangle L., Laguncularia racemosa Gaert and Avicennia germinans (L.)L. Experiments were performed with adult plants growing naturally in the field under a salinity of 35‰. Gas exchange parameters showed that A. germinans had significantly higher photosynthetic rates, and lower stomatal conductance and transpiration rates, compared to the other two mangroves. In concert with this, instantaneous water use efficiency was significantly high in A. germinans, intermediate in L. racemosa and lowest in R. mangle. The hydraulic parameters of the three mangrove species were in the lowest end of the range reported for tropical trees. However, the three mangrove species exhibited measurable differences in hydraulic parameters related to the control of water requirements for maintenance of carbon gain. L. racemosa and A. germinans showed less efficient water transport at shoot level but were the more efficient species in water use at the leaf level in comparison to R. mangle. Received: 7 April 1999 / Accepted: 25 July 1999     

Water relations of the root hemiparasite Olax phyllanthi (Labill) R.Br. (Olacaceae) and its multiple hosts

Summary Water relations of the root hemiparasite Olax phyllanthi were compared with those of its major species of hosts in natural habitat in coastal heath near Denmark, SW Australia. Leaf water potentials of Olax during winter were 0.4 to 1.4 MPa lower (more negative) than those of all (29) non parasitic host species examined. During the dry summer months (January to March), shallow-rooted hosts developed water potentials up to 3 MPa lower than those of Olax, and were accordingly rated as no longer accessible as a source of water to the hemiparasite. By contrast, deep-rooted hosts, with access to the water table, showed water potentials less negative than Olax, and haustorial contacts retained with these apparently enabled continued extraction of water and nutrients throughout the summer. Three other species of root hemiparasites parasitized by Olax, but not themselves parasitizing Olax, showed leaf water potentials throughout the year very close to, and mostly slightly more negative than those of Olax. Nocturnal measurements of leaf water potential in winter (July and August) in soil at field capacity (water potential –0.006 MPa) showed maintenance of a 0.5–0.8 MPa potential difference between Olax and a range of common host species. By dawn most hosts had equilibrated with the water potential of the soil, whereas both exposed and bagged Olax plants recorded potentials of –0.8 MPa. Daytime rates of transpiration and photosynthesis of Olax were less than those of a range of common hosts, but water use efficiencies were not consistently different between hemiparasite and hosts. This was reflected in almost identical mean values for carbon isotope ratio (¹³C/¹²C) between Olax (mean value –27.0) and thirteen frequently exploited hosts ( value –27.1). The results are discussed in relation to published information on other angiosperm hemiparasites.     

Carbon exchange and water use efficiency of a growing, irrigated olive orchard

We measured eddy covariance fluxes of CO₂ and H₂O over a flat irrigated olive orchard during growth, in different periods from Leaf Area Index (LAI) of 0.3–1.9; measurements of soil respiration were also collected. The daily net ecosystem exchange flux (F_NEE) was practically zero at LAI around 0.4 or when the orchard intercepted 11% of the incoming daily radiation; at the end of the experiment, with LAI of 1.9 (and the fraction of intercepted daily radiation close to 0.5), F_NEE was around 10 g CO₂ m⁻² day⁻¹. The night-time ecosystem respiration (R_eco), calculated from eddy fluxes in well-mixed night conditions, show a clear but non-linear dependence with LAI; it ranged from 0.05 to 0.15 mg CO₂ m⁻² s⁻¹ (in average), being the lower limit ideally close to the heterotrophic soil respiration at the site. The gross primary production flux (F_GPP) was linearly related to LAI within the LAI range of this experiment (with 11 g CO₂ m⁻² day⁻¹ increments per unit of LAI) and to the fraction of intercepted radiation. The maximum rates of F_GPP (0.75 mg CO₂ m⁻² s⁻¹) were obtained in the summer mornings of 2002, at LAI close to 1.9. F_GPP was strongly modulated by vapour pressure deficit (VPD) through the canopy conductance, even in absence of water stress. Hence, especially in the summer, the maximum rates of carbon assimilation are reached always before noon. The daily course of F_GPP shows a two-phase pattern, first related to irradiance and then to canopy conductance. The water use efficiency (WUE) was, in average, 3.8, 6.3 and 7 g CO₂ L⁻¹ in 1999, 2001 and 2002, respectively, with maxima always in the early morning. Hourly WUE was strongly related to VPD (WUE = −10.25 + 22.52 × VPD^−0.34). Our results suggest that drip irrigated orchards in general, and olive in particular, deserve specific carbon exchange and carbon budget studies and cannot be easily included in other biomes.