Carbon assimilation, nitrogen, and photochemical efficiency of different Himalayan tree species along an altitudinal gradient

In the area of Jumla region in Western Nepal, measurements of saturated leaf net photosynthetic rate (P_sat), nitrogen content, leaf fluorescence, carbon isotopic composition, and water status were performed on woody coniferous (Pinus wallichiana, Picea smithiana, Abies spectabilis, Juniperus wallichiana, Taxus baccata), evergreen (Quercus semecarpifolia, Rhododendron campanulatum), and deciduous broadleaved species (Betula utilis, Populus ciliata, Sorbus cuspidata) spreading from 2 400 m up to the treeline at 4 200 m a.s.l. With the exception of J. wallichiana, P_sat values were lower in coniferous than broadleaved species. Q. semecarpifolia, that in this area grows above the coniferous belt between 3 000 and 4 000 m, showed the highest P_sat at saturating irradiance and the highest leaf N content. This N content was higher and P_sat lower than those of evergreen oak species of tempe forests at middle and low altitudes. For all species, P_sat and N content were linearly correlated, but instantaneous nitrogen use efficiency was lower than values measured in lowland and temperate plant communities. The values of carbon isotopic composition, estimated by ₁₃C, showed the same range reported for temperate tree species. The ranking of ₁₃C values for the different tree types was conifers < evergreen broadleaved13C were found along the altitudinal gradient. Quantum yield of photochemistry at saturating irradiance, measured by leaf fluorescence (F/F_m), was highest in J. wallichiana and lowest in T. baccata. Overall, photochemical efficiency was more strongly related to species than to altitude. Interestingly, changes of .F/F_m along the altitudinal gradient correlated well with the reported altitudinal distribution of the species.This revised version was published online in March 2005 with corrections to the page numbers.     

Adaptive photosynthetic strategies of the Mediterranean maquis species according to their origin

In consideration of their origin the adaptive strategies of the evergreen species of the Mediterranean maquis were analysed. Rosmarinus officinalis L., Erica arborea L., and Erica multiflora L. had the lowest net photosynthetic rate (P_N) in the favourable period [7.8±0.6 μmol(CO₂) m^?2s^?1, mean value], the highest P_N decrease (on an average 86 % of the maximum) but the highest recovery capacity (>70 % of the maximum) at the first rainfall in September. Cistus incanus L. and Arbutus unedo L. had the highest P_N during the favourable period [15.5±5.2 μmol(CO₂) m^?2s^?1, mean value], 79 % decrease during drought, and a lower recovery capacity (on an average 54 %). Quercus ilex L., Phillyrea latifolia L., and Pistacia lentiscus L. had an intermediate P_N in the favourable period [9.2±1.3 μmol(CO2) m^?2s^?1, mean value], a lower reduction during drought (on an average 63 %), and a range from 62 % (Q. ilex and P. latifolia) to 39 % (P. lentiscus) of recovery capacity. The Mediterranean species had higher decrease in P_N and stomatal conductance during drought and a higher recovery capacity than the pre-Mediterranean species. Among the pre-Mediterranean species, P. latifoliahad the best adaptation to long drought periods also by its higher leaf mass per area (LMA) which lowered leaf temperature thus decreasing transpiration rate during drought. Moreover, its leaf longevity determined a more stable leaf biomass during the year. Among the Mediteranean species, R. officinalis was the best adapted species to short drought periods by its ability to rapidly recover. Nevertheless, R. officinalis had the lowest tolerance to high temperatures by its P_N dropping below half its maximum value when leaf temperature was over 33.6°C. R. officinalismay be used as a bioindicator species of global change.     

The effects of vegetation types and microhabitats on carabid beetle community composition in cool temperate Japan

The effects of vegetation types and environmental factors on carabid beetle (Coleoptera: Carabidae) communities were studied. Carabid beetles were collected using pitfall traps (total 2844 trapping days) and seven microenvironmental factors were measured in four vegetation types: grassland, natural evergreen coniferous forest (Pinus densiflora), deciduous broad-leaved natural forest (Quercus crispula, Betula platyphylla, Alnus japonica, or Fagus crenata), and deciduous coniferous plantation (Larix kaempferi) in cool temperate Japan. These four vegetation types provided a novel comparison between natural forests and plantations because the vast majority of related studies have investigated only deciduous broad-leaved natural forests and evergreen coniferous plantations. PERMANOVA indicated that vegetation types affected carabid community composition. Ordination plots showed that community composition differed greatly between grassland and forest vegetation types, but that community composition in the plantation forest overlapped with that of natural forest types. Characteristics differentiating the grassland included a high proportion of winged species and a low mean carabid body weight. Among the examined environmental factors, litter depth, soil water content, and depth of the soil A-horizon had large effects on carabid communities. These results suggest that the effect of afforestation on carabid communities in cool temperate Japan might be insignificant compared with the effects of cover types (deciduous vs. evergreen) and microenvironmental factors.     

Leaf age dependent changes in within-canopy variation in leaf functional traits: a meta-analysis

Within-canopy variation in leaf structural and photosynthetic characteristics is a major means by which whole canopy photosynthesis is maximized at given total canopy nitrogen. As key acclimatory modifications, leaf nitrogen content (N _A) and photosynthetic capacity (A _A) per unit area increase with increasing light availability in the canopy and these increases are associated with increases in leaf dry mass per unit area (M _A) and/or nitrogen content per dry mass and/or allocation. However, leaf functional characteristics change with increasing leaf age during leaf development and aging, but the importance of these alterations for within-canopy trait gradients is unknown. I conducted a meta-analysis based on 71 canopies that were sampled at different time periods or, in evergreens, included measurements for different-aged leaves to understand how within-canopy variations in leaf traits (trait plasticity) depend on leaf age. The analysis demonstrated that in evergreen woody species, M _A and N _A plasticity decreased with increasing leaf age, but the change in A _A plasticity was less suggesting a certain re-acclimation of A _A to altered light. In deciduous woody species, M _A and N _A gradients in flush-type species increased during leaf development and were almost invariable through the rest of the season, while in continuously leaf-forming species, the trait gradients increased constantly with increasing leaf age. In forbs, N _A plasticity increased, while in grasses, N _A plasticity decreased with increasing leaf age, reflecting life form differences in age-dependent changes in light availability and in nitrogen resorption for growth of generative organs. Although more work is needed to improve the coverage of age-dependent plasticity changes in some plant life forms, I argue that the age-dependent variation in trait plasticity uncovered in this study is large enough to warrant incorporation in simulations of canopy photosynthesis through the growing period.     

Influence of phosphorus and nitrogen on photosynthetic parameters and growth in <Emphasis Type="Italic">Vicia faba</Emphasis> L.

The influence of phosphorus (P) and nitrogen (N) supply on biomass, leaf area, photon saturated photosynthetic rate (P_max), quantum yield efficiency (α), intercellular CO₂ concentration (C_i), and carboxylation efficiency (CE) was investigated in Vicia faba. The influence of P on N accumulation, biomass, and leaf area production was also investigated. An increase in P supply was consistently associated with an increase in N accumulation and N productivity in terms of biomass and leaf area production. Furthermore, P increased the photosynthetic N use efficiency (NUE) in terms of P_max and α. An increase in P supply was also associated with an increase in CE and a decrease in C_i. Under variable daily meteorological conditions specific leaf nitrogen content (N_L), specific leaf phosphorus content (P_L), specific leaf area (δ_L), root mass fraction (R_f), P_max, and α remained constant for a given N and P supply. A monotonic decline in the steady-state value of R_f occurred with increasing N supply. δ_L increased with increasing N supply or with increasing N_L. We tested also the hypothesis that P supply positively affects both N demand and photosynthetic NUE by influencing the upper limit of the asymptotic values for P_max and CE, and the lower limit for C_i in response to increasing N.     

Early <Emphasis Type="Italic">Acacia</Emphasis> invasion in a sandy ecosystem enables shading mediated by soil,leaf nitrogen and facilitation

Australian species of the genus Acacia are amongst the most invasive trees. As nitrogen fixers, they are able to invade oligotrophic ecosystems and alter ecosystem functioning to their benefit. We aimed to answer three questions: How does early Acacia invasion influence nitrogen and light in a sandy savanna? How does early Acacia invasion impact biodiversity? Does early invasion alter ecosystem functioning towards the dominance of Acacia? We analyzed (using generalized linear mixed models and richness estimators) paired plots focused on plants of Acacia mangium (Fabaceae) and plants of Marcetia taxifolia (Melastomataceae) by taking hemispherical photos and sampling plants, leaves and soil for measurements of light, richness, leaf nitrogen, leaf δ15N, soil nitrogen and soil coarse sand. The results suggest that early Acacia invasion alters the control of soil and of leaf nitrogen and increases shading, enabling a much wider range of light variation. The δ15N results suggest that the nitrogen taken up by Acacia is transferred to neighboring plants and influences the light environment, suggesting facilitation. The enrichment of plant species observed during early Acacia invasion is consistent with the wider range of light variation, but the forecasted leaf nitrogen conditions during the established phase of Acacia invasion might cause loss of light-demanding species because of increased shading. If early Acacia invasion turns into an established phase with highly increased shading, Acacia seedlings might be favored and ecosystem functioning might change towards its dominance.     

Gas exchanges of an endangered species <Emphasis Type="Italic">Syringa pinnatifolia</Emphasis> and a widespread congener <Emphasis Type="Italic">S. oblata</Emphasis>

Net photosynthetic rate (P_N), transpiration rate (E), water use efficiency (WUE), stomatal conductance (g_s), and stomatal limitation (L_s) were investigated in two Syringa species. The saturation irradiance (SI) was 400 µmol m^-2s^-1 for S. pinnatifolia and 1 700 µmol m^-2s^-1 for S. oblata. Compared with S. oblata, S. pinnatifolia had extremely low g _s . Unlike S. oblata, the maximal photosynthetic rate (P_max) in S. pinnatifoliaoccurred around 08:00 and then fell down, indicating this species was sensitive to higher temperature and high photosynthetic photon flux density. However, such phenomenon was interrupted by the leaf development rhythms before summer. A relatively lower P_N together with a lower leaf area and shoot growth showed the capacity for carbon assimilation was poorer in S. pinnatifolia.     

Solute patterns and diurnal variation of photosynthesis and chlorophyll fluorescence in Korean coastal sand dune plants

Four plant species, Elymus mollis Trin., Carex kobomugi Ohwi, Glehnia littoralis F. Schmidt ex Miq., and Vitex rotundifolia L.f., are dominant perennial species in coastal sand dunes of Korea. We examined a physiological adaptation of these species by measurements of diurnal variation in photosynthesis and chlorophyll (Chl) fluorescence and solute patterns in leaves during one season (June), which is favorable for plant growth of all four species. All four species adopted different strategies in order to utilize radiation and to maintain water status under a fluctuating microclimate. Although the lowest water contents among four plant species was found, E. mollis with a high Chl and K⁺ content showed better photosynthetic performance, with high stomatal conductance (g _s), net photosynthetic rate (P _N), instantaneous carboxylation efficiency (CE), and water-use efficiency. Midday depression of P _N in E. mollis and G. littoralis, without a reduction of gs, was associated with a reduction in CE and maximum photochemical efficiency of PSII, indicating nonstomatal limitation. Photosynthesis depression in both C. kobomugi and V. rotundifolia, with relatively low g _s values, could be attributed to both stomatal and nonstomatal limitations. The high storage capacity for inorganic ions in E. molli, C. kobomugi, and G. littoralis may play an efficient role in regulating photosynthesis and maintaining leaf water status through stomatal control, and can also play an important role in osmotic adjustment.     

The coupling effects of water deficit and nitrogen supply on photosynthesis,WUE, and stable isotope composition in <Emphasis Type="Italic">Picea asperata</Emphasis>

Water stress and nitrogen (N) availability are the two main factors limiting plant growth, and the two constrains can interact in intricate ways. Moreover, atmospheric N depositions are altering the availability of these limiting factors in many terrestrial ecosystems. Here, we studied the combined effects of different soil water availability and N supply on photosynthesis and water-use efficiency (WUE) in Picea asperata seedlings cultured in pots, using gas exchange, and stable carbon and nitrogen isotope composition (δ ¹³C and δ ¹⁵N). Photosynthesis under light saturation (A _sat) and stomatal conductance (g _s) of P. asperata decreased as the soil moisture gradually diminished. Under severe water-stress condition, N addition decreased the A _sat and g _s, whereas the positive effects were observed in moderate water-stress and well-watered conditions. The effect of N addition on the intrinsic WUE (WUE_i) deduced from gas exchange was associated with soil water availability, whereas long-term WUE evaluated by leaf δ ¹³C only affected by soil water availability, and it would be elevated with soil moisture gradually diminished. Water deficit would restrict the uptake and further transport of N to the aboveground parts of P. asperata, and then increasing δ ¹⁵N. Therefore, δ ¹⁵N in plant tissues may reflect changes in N allocation within plants. These results indicate that the effect of N enrichment on photosynthesis in P. asperata is largely, if not entirely, dependent on the severity of water stress, and P. asperata would be more sensitive to increasing N enrichment under low soil water availability than under high soil moisture.     

Ecophysiological responses of native invasive woody <Emphasis Type="Italic">Juniperus virginiana</Emphasis> L. to resource availability and stand characteristics in the semiarid grasslands of the Nebraska Sandhills

Vegetation in grasslands is changing at an unprecedented rate. In the Nebraska Sandhills, this shift is attributed in part to encroachment of the woody species Juniperus virginiana. We investigated changes in resource availability and their feedback on seasonal trends in photosynthetic characteristics of J. virginiana trees scattered in open grasslands vs. a dense 57-year-old stand. Dense stand exhibited lower volumetric soil water content, NH₄ ⁺, NO₃ ^–, and δ¹³C, as well as foliage δ¹³C, δ¹⁵N, and N content, compared to grasslands. Water potential was higher in trees in grasslands compared to dense stand. J. virginiana in dense stand exhibited similar trends to trees in grasslands for net photosynthetic rate (P _N), stomatal conductance, transpiration, maximum photochemical efficiency of PSII, maximum carboxylation velocity, and maximum rate of electron transport. P _N peaked early summer and declined in the fall, with trees in open grasslands lagging behind those in dense stand. Plasticity of this species may place it at a competitive advantage in the Sandhills, further altering grasslands vegetation and ecosystem processes.     

Low assimilation efficiency of photorespiratory ammonia in conifer leaves

Glutamine synthetase (GS) localized in the chloroplasts, GS2, is a key enzyme in the assimilation of ammonia (NH₃) produced from the photorespiration pathway in angiosperms, but it is absent from some coniferous species belonging to Pinaceae such as Pinus. We examined whether the absence of GS2 is common in conifers (Pinidae) and also addressed the question of whether assimilation efficiency of photorespiratory NH₃ differs between conifers that may potentially lack GS2 and angiosperms. Search of the expressed sequence tag database of Cryptomeria japonica, a conifer in Cupressaceae, and immunoblotting analyses of leaf GS proteins of 13 species from all family members in Pinidae revealed that all tested conifers exhibited only GS1 isoforms. We compared leaf NH₃ compensation point (γ_NH3) and the increments in leaf ammonium content per unit photorespiratory activity (NH₃ leakiness), i.e. inverse measures of the assimilation efficiency, between conifers (C. japonica and Pinus densiflora) and angiosperms (Phaseolus vulgaris and two Populus species). Both γ_NH3 and NH₃ leakiness were higher in the two conifers than in the three angiosperms tested. Thus, we concluded that the absence of GS2 is common in conifers, and assimilation efficiency of photorespiratory NH₃ is intrinsically lower in conifer leaves than in angiosperm leaves. These results imply that acquisition of GS2 in land plants is an adaptive mechanism for efficient NH₃ assimilation under photorespiratory environments.     

Evolution of protein transport to the chloroplast envelope membranes

Comparing with other angiosperms, most members within the family Orchidaceae have lower photosynthetic capacities. However, the underlying mechanisms remain unclear. Cypripedium and Paphiopedilum are closely related phylogenetically in Orchidaceae, but their photosynthetic performances are different. We explored the roles of internal anatomy and diffusional conductance in determining photosynthesis in three Cypripedium and three Paphiopedilum species, and quantitatively analyzed their diffusional and biochemical limitations to photosynthesis. Paphiopedilum species showed lower light-saturated photosynthetic rate (A _N), stomatal conductance (g _s), and mesophyll conductance (g _m) than Cypripedium species. A _N was positively correlated with g _s and g _m. And yet, in both species A _N was more strongly limited by g _m than by biochemical factors or g _s. The greater g _s of Cypripedium was mainly affected by larger stomatal apparatus area and smaller pore depth, while the less g _m of Paphiopedilum was determined by the reduced surface area of mesophyll cells and chloroplasts exposed to intercellular airspace per unit of leaf area, and much thicker cell wall thickness. These results suggest that leaf anatomical structure is the key factor affecting g _m, which is largely responsible for the difference in photosynthetic capacity between those two genera. Our findings provide new insight into the photosynthetic physiology and functional diversification of orchids.     

Rearrangement of leaf traits with changing source-sink relationship in blueberry (Vaccinium corymbosum L.) leaves

The source-sink relationship is one of major determinants of plant performance. The influence of reproductive sink demand on light-saturated photosynthesis (P_max), dark respiration (R_D), stomatal conductance (g_s), intrinsic water-use efficiency (WUE_i), contents of soluble sugar (SSC), nitrogen, carbon, and photosynthetic pigments was examined in blueberry (Vaccinium corymbosum L. cv. ‘Brigitta’) during the final stage of rapid fruit growth. Measurements were performed three times per day on developed, sun-exposed leaves of girdled shoots with 0.1, 1, and 10 fruit per leaf (0.1F:L, 1F:L, and 10F:L, respectively) and nongirdled shoots bearing one fruit per leaf (NG). Girdling and lower fruit amount induced lower P_max, g_s, N, and total chlorophyll (Chl) and higher WUE_i, SSC, R_D, Chl a/b ratio and carotenoids-to-chlorophylls ratio (Car/Chl) for the 1F:L and 0.1F:L treatments. The impact of girdling was counterbalanced by 10F:L, with NG and 10F:L having similar values. Variables other than P_max, R_D, g_s, WUE_i, and SSC were unaffected throughout the course of the day. P_max and g_s decreased during the course of the day, but g_s decreased more than P_max in the afternoon, while WUE_i was increasing in almost all treatments. SSC increased from the morning until afternoon, whereas R_D peaked at noon regardless of the treatment. Generally, P_max was closely and negatively correlated to SSC, indicating that sugar-sensing mechanisms played an important role in regulation of blueberry leaf photosynthesis. With respect to treatments, P_max and N content were positively related, while R_D was not associated to substrate availability. The enhanced Car/Chl ratio showed a higher photoprotection under the lower sink demand. Changes in the source-sink relationship in ‘Brigitta’ blueberry led to a rearrangement of physiological and structural leaf traits which allowed adjusting the daily balance between carbon assimilation and absorbed light energy.     

Morphological and molecular investigations of <Emphasis Type="Italic">Gagea</Emphasis> (Liliaceae) in southeastern Kazakhstan with special reference to putative altitudinal hybrid zones

The most important center of speciation in the genus Gagea is thought to be in Central Asia. Here, we focus on species diversity in southeastern Kazakhstan (around Almaty, Ili-Alatau range of the Western Tian-Shan mountains). We studied an elevational transect, reaching from lowland steppes to the alpine zone (500–2750 m a. s. l.), and carried out detailed morphological and molecular investigations for populations of Gagea spp. Nine species were identified in different altitudinal zones; one of these (Gagea almaatensis) is described as new to science. We could detect two altitudinal contact zones between closely related species: G. filiformis and G. granulosa (sect. Minimae), and G. almaatensis and G. kuraminica (sect. Gagea). Morphological and molecular investigations (ITS data and cpDNA networks) indicate ongoing hybridization of co-occurring G. filiformis into G. granulosa and putative bidirectional hybridization events between G. almaatensis and G. kuraminica.     

Parameters of photosynthesis light curve in <Emphasis Type="Italic">Salix dasyclados</Emphasis> and their changes during the growth season

A dependence of the photosynthesis rate on light is characterized by a number of parameters that are often used for comparison between plant species or for finding photosynthesis interconnections with other physiological processes. In order to properly assessed these parameters, we measured the maximum apparent photosynthesis rate (P _max), dark respiration rate (R _d), light compensation point (LCP), quantum yield corresponding to photosynthetic efficiency (QY), and the light saturation constant (K _s), taking into consideration the leaf plastochron index during vegetation of one of the willow species (Salix dasyclados Wimn.). The P _max value was the highest in the beginning of the growth season when the leaf reached 65% of its full area; after that P _max slowly declined. The most important cardinal value for R _d is its plateau reached by the end of leaf growth, i.e., later than the photosynthesis rate maximum. This plateau value also decreased during vegetation. The LCP value changed in the same way as R _d but reached its plateau simultaneously with the photosynthesis rate maximum. QY also reached its maximum at the same time with the photosynthesis rate; during vegetation it changed more than twofold. The K _s value also changed almost twofold during the season, reaching its maximum together or slightly later than the photosynthesis maximum and then remained constant. Thus, we have found significant changes in the parameters of the photosynthesis light curve during growth season. This shows that they can be used only after a thorough study of leaf development in each particular plant species. Usually performed measuring gas exchange parameters in fully developed leaves does not yield their maximum values and thus does not have any physiological sense.     

Mitochondrial DNA sequence variation in Drosophilid species (Diptera: Drosophilidae) along altitudinal gradient from Central Himalayan region of India

Central Himalayan region of India encompasses varied ecological habitats ranging from near tropics to the mid-elevation forests dominated by cool-temperate taxa. In past, we have reported several new records and novel species from Uttarakhand state of India. Here, we assessed genetic variations in three mitochondrial genes, namely, 16S rRNA, cytochrome c oxidase subunit I and cytochrome c oxidase subunit II (COI and COII) in 26 drosophilid species collected along altitudinal transect from 550 to 2700 m above mean sea level. In the present study, overall 543 sequences were generated, 82 for 16S rRNA, 238 for COI, 223 for COII with 21, 47 and 45 mitochondrial haplotypes for 16S rRNA, COI and COII genes, respectively. Almost all species were represented by 2–3 unique mitochondrial haplotypes, depicting a significant impact of environmental heterogeneity along altitudinal gradient on genetic diversity. Also for the first time, molecular data of some rare species like Drosophila mukteshwarensis, Liodrosophila nitida, Lordiphosa parantillaria, Lordiphosa ayarpathaensis, Scaptomyza himalayana, Scaptomyza tistai, Zaprionus grandis and Stegana minuta are provided to public domains through this study.     

Stomatal regulation in Douglas fir following a fungal-mediated chronic reduction in leaf area

Pathogens can cause chronic premature needle abscission in coniferous species. To assess the potential impacts on tree productivity, stomatal regulation was investigated in Douglas fir with chronic stomatal occlusion and defoliation from varying levels of the Swiss needle cast (SNC) fungus, Phaeocryptopus gaeumannii. Levels of SNC disease and subsequent defoliation were manipulated by choosing six sites with varying levels of disease and by foliar applications of fungicides on six trees per site. Diurnal measurements of leaf water potential (Ψ_leaf), stomatal conductance (g _s) and vapor pressure deficit (D) were made on six fungicide treated and six control trees per site. In addition, leaf specific hydraulic conductance was calculated on a single branch (K _{L_B}) from three trees per treatment per site. Stomatal conductance at D=1 kPa (g _sref) was negatively correlated with fungal colonization (number of fruiting bodies present in needle stomata) and positively correlated with K _{L_B}. Despite reduced needle retention in diseased trees, K _L declined due to a reduction in sapwood area and permeability (i.e., increasing presence of latewood in functional sapwood). In general, stomatal sensitivity to D for all foliage was consistent with stomatal regulation based on a simple hydraulic model [g _s=K _L(Ψ_soil?Ψ_leaf)/ D], which assumes strict stomatal regulation of Ψ_leaf. However, when fungal presence reduced maximum g _s below the potential maximum supported by hydraulic architecture, stomatal sensitivity was lower than expected based on the theoretical relationship: dg _s/dlnD=0.6·g _sref. The results indicate that losses in productivity associated with physical blockage of stomata and defoliation are compounded by additional losses in K _L and a reduction in g _s in remaining functional stomata.     

Light availability and soil flooding regulate photosynthesis of an imperiled shrub in lowland forests of the Mississippi Alluvial Valley,USA

Physiological responses to light availability and soil flooding on Lindera melissifolia (Walt.) Blume were studied. Shrubs were grown under 70, 37 or 5% of full sunlight with either 0, 45, or 90 d of soil flooding. We measured leaf photosynthetic rate (PN) to test the hypothesis that soil flooding reduces P _N in L. melissifolia following shrub acclimation to low light availability. Results showed that light availability and soil flooding interacted to affect P _N. In the 0 d and 45 d flooding regimes (flood water removed 36–39 d prior to measurement), P _N was similar between shrubs receiving 70% or 37% light, and these shrubs had 147% greater P _N than shrubs receiving 5% light. Shrubs receiving 90 d of soil flooding had similar low rates of area-based P _N regardless of light level. Similar P _N between 0 d and 45 d flooded shrubs indicated physiological recovery following removal of flood water.