Spectral changes with leaf aging in Amazon caatinga

 Significant gaps exist in the knowledge of tropical leaf spectra and the manner in which spectra change as leaves age in their natural environment. Leaf aging effects may be particularly important in tropical vegetation growing on nutrient poor soils, such as Amazon caatinga, a white sand community common in the Amazon Basin. Spectral changes observed in six caatinga dominants include decreased reflectance and transmittance and increased absorptance for epiphyll-coated older leaves. Near-infrared (NIR) changes were most significant. More detailed spectral and physical changes were studied in one dominant, Aldina heterophylla. Over 16 months, Aldina study plants produced one or two leaf flushes. During leaf expansion, leaf water content and Specific Leaf Area decreased rapidly. Over the first 6 months spectral changes occurred across the spectrum, resulting in decreased transmittance and increased absorptance in the visible and NIR and decreased visible and increased NIR reflectance. In contrast, significant spectral changes were restricted to the NIR over the last 9 months, which showed a 10% absorptance increase associated primarily with increasing epiphylls and necrosis. At the canopy scale, increased NIR absorptance provides a mechanism for producing seasonally varying forest albedo and changing NIR to red ratios, independent of changes in other canopy attributes. In the Amazon caatinga studied, all canopy dominants were subject to epiphyllic growth providing a mechanism for distinguishing these forest types spectrally from more diverse terra-firme forest or forest types with more rapid leaf turnover, such as second growth. These changes are observable using remote sensing and could be used to map caatinga and monitor interannual or seasonal variability in phenology. If these results can be extended to other communities with long-lived foliage, they may offer a means for mapping vegetation on the basis of leaf longevity. Received: 18 November 1996/Accepted: 24 December 1997     

Population- and family-level variation of brittlebush (Encelia farinosa, Asteraceae) pubescence: its relation to drought and implications for selection in variable environments

Because leaf pubescence of the desert shrub Encelia farinosa increases in response to drought and influences photosynthesis and transpiration, we hypothesized that differences in water availability across the range of this species may result in genetic differentiation for pubescence and associated productivity traits. We examined maternal family variation of pubescence-moderated light absorption (absorptance) in three populations of E. farinosa. Absorptance was always greatest for plants from the high-rainfall environment and lowest for those from the driest site, but the rate of absorptance change in response to drought was similar among all populations. Similar patterns were found when we compared families within populations-all genotypes had similar initial leaf absorptances, differentiated very early in seasonal growth, then had concordant changes in absorptance thereafter. However, family-level variance was greatest for plants from the driest site, a region with highly heterogeneous precipitation patterns, whereas low variance was found for plants from the wettest, least heterogeneous site. The concordance of leaf absorptance changes, within and among populations, may be due to integration with other drought-related traits; however, the differences in absorptance values within and among populations suggest that variation of leaf pubescence results from selection associated with geographical and local patterns of water availability.     

Variation in leaf longevity of Pistacia lentiscus and its relationship to sex and drought stress inferred from leaf δ13C

1. Leaf formation, loss, retention, longevity and biomass on male branches of the evergreen mediterranean shrub Pistacia lentiscus , L. correlated strongly with water-use efficiency inferred from leaf δ¹³C across a gradient of precipitation on the island of Mallorca, Spain.
2. The correlations suggest that the leaf phenology is under control of drought-induced constraints on the carbon balance.
3. In fruiting female branches, the correlations between the inferred water-use efficiency and number of formed and retained leaves, leaf biomass and leaf longevity were non-significant. Leaf formation was strongly reduced by fruiting and the females compensated the reduced photosynthetic capacity by retaining older leaves for a longer time than male plants.
4. It is suggested that leaf longevity in females is under strong control of resource allocation to fruit formation which is 'overlaid' on the drought-induced carbon stress, which led to the observed longer leaf longevity in females than in males.     

THE INFLUENCE OF WATER STRESS AND TEMPERATURE ON LEAF PUBESCENCE DEVELOPMENT IN ENCELIA FARINOSA

The degree of leaf pubescence development in the arid land shrub Encelia farinosa Gray is affected by air temperature, leaf water potential, and previous history of the apical meristem during the current growing season. Changes in leaf pubescence levels change leaf spectral characteristics and affect both leaf temperature and photosynthesis. Decreasing leaf water potentials and increasing air temperatures both independently increase pubescence development as measured by decreased leaf absorptances. During any one growing season leaf absorptance may change reversibly coincident with air temperature changes, but with respect to water stress leaf absorptance only decreases as the season progresses. The ecological significance of regulation of the leaf spectral characteristics is discussed.     

Pubescence and leaf spectral characteristics in a desert shrub,Encelia farinosa

Summary The effects of leaf hairs (pubescence) on leaf spectral characteristics were measured for the drought-deciduous desert shrub Encelia farinosa. Leaf absorptance to solar radiation is diminished by the presence of pubescence. The pubescence appears to be reflective only after the hairs have dried out. There are seasonal changes in leaf absorptance; leaves produced at the beginning of a growing season have high absorptances, whereas leaves produced during the growing season are more pubescent and have lower absorptances. The decrease in leaf absorptance is the result of an increase in pubescence density and thickness. Between 400 and 700 nm (visible wavelengths), pubescence serves as a blanket reflector. However, over the entire solar spectrum (400–3000 nm), the pubescence preferentially reflects near infrared radiation (700–3000 nm) over photosynthetically useful solar radiation (400–700 nm). Leaf absorptance to solar radiation (400–3000 nm) varies between 46 and 16%, depending on pubescence; whereas leaf absorptance to photosynthetically useful radiation (400–700 nm) may vary from 81 to 29%.C.I.W.-D.P.B. Publication No. 612     

Genotypic variation in drought response of silver birch (Betula pendula Roth): leaf and root morphology and carbon partitioning

This study investigates the drought response of four genotypes of Betula pendula with a focus on leaf and root morphological traits, leaf phenology and carbon partitioning between shoot and root. Potted one-year-old clonal plants of four genotypes from regions with low to high annual rainfall (550–1270 mm year⁻¹) were subjected to drought periods of 12–14 weeks in two subsequent years. Well-watered control plants of the four genotypes differed significantly with respect to total leaf area per plant (LA) and specific leaf area (SLA), whereas differences in total fine root surface area (RA), root specific area (SRA), and the fine root:leaf mass ratio (FR:LM) were not significant. Highest LA and SLA were found in the clone originating from the driest environment. In complementary physiological investigations this clone was found to have the highest water use as well which was interpreted as competitive superiority in terms of water consumption. Drought resulted in an increase in SLA in all genotypes, and a decrease in LA. Leaf area reduction was more pronounced in the genotypes from high than in those from low rainfall origin. The ratio of total root to leaf surfaces remained more or less constant after drought application despite an increase in FR:LM. This is explained by a decrease in SRA resulting from a reduced abundance of very small fine rootlets (diameter <0.2 mm) in the drought-treated plants. The loss in total root surface area due to a reduction in finest root mass was compensated for by a relative increase in total root dry mass per plant. Comparison of results from the first and second drought period indicated a marked influence of timing of drought, root system size, and putative root limitation on plant drought response. We conclude that leaf and root morphology, the total leaf and root surfaces, and the morphological response to drought in birch are to a large extent under genetic control.     

Deciduous leaf drop reduces insect herbivory

Deciduous leaf fall is thought to be an adaptation that allows plants living in seasonal environments to reduce water loss and damage during unfavorable periods while increasing photosynthetic rates during favorable periods. Observations of natural variation in leaf shedding suggest that deciduous leaf fall may also allow plants to reduce herbivory. I tested this hypothesis by experimentally manipulating leaf retention for Quercus lobata and observing natural rates of herbivory. Quercus lobata is primarily deciduous although individuals show considerable natural variation in leaf retention. Oak saplings with no leaves through winter experienced reduced attack by cynipid gall makers the following spring. This pattern was consistent with the positive correlation between natural leaf persistence and gall numbers. These cynipids do not overwinter on the leaves that trees retain through winter, although they appear to use persistent leaves as oviposition cues. If these results are general for woody plants in continental temperate habitats, they suggest that an important and unrecognized consequence of deciduous leaf shedding may be a reduction in herbivore damage, and that this effect should be included in models of deciduous and evergreen behavior.     

Leaf physiology does not predict leaf habit; examples from tropical dry forest

Leaf structure and physiology are thought to be closely linked to leaf longevity and leaf habit. Here we compare the seasonal variation in leaf hydraulic conductance (k_leaf) and water potential of two evergreen tree species with contrasting leaf life spans, and two species with similar leaf longevity but contrasting leaf habit, one being deciduous and the other evergreen. One of the evergreen species, Simarouba glauca, produced relatively short-lived leaves that maintained high hydraulic conductance year round by periodic flushing. The other evergreen species, Quercus oleoides, produced longer-lived leaves with lower k_leaf and as a result minimum leaf water potential was much lower than in S. glauca (–2.8 MPa vs –1.6 MPa). Associated with exposure to lower water potentials, Q. oleoides leaves were harder, had a higher modulus of elasticity, and were less vulnerable to cavitation than S. glauca leaves. Both species operate at water potentials capable of inducing 20 (S. glauca) to 50% (Q. oleoides) loss of k_leaf during the dry season although no evidence of cumulative losses in k_leaf were observed in either species suggesting regular repair of embolisms. Leaf longevity in the deciduous species Rhedera trinervis is similar to that of S. glauca, although maximum k_leaf was lower. Furthermore, a decline in leaf water potential at the onset of the dry season led to cumulative losses in k_leaf in R. trinervis that culminated in leaf shedding.     

Characteristics of Encelia species differing in leaf reflectance and transpiration rate under common garden conditions

Summary The performance of coastal and desert species of Encelia (Asteraceae) were evaluated through common garden growth observations. The obectives of the study were to evaluate the roles of leaf features, thought to be of adaptive value (increased leaf reflectance and/or transpirational cooling), on plant growth in the hot, arid, desert garden versus their impact on growth under cooler, relatively more moist coastal garden conditions. E. californica native to the coast of southern California and E. farinosa, and E. frutescens, interior desert species, were grown in common gardens at coastal (Irvine, California) and interior (Phoenix, Arizona) sites under both irrigated and natural conditions. Although all species survived in both gardens during the two and a half year study period, there were large differences in their sizes. In the desert garden, leaf conductance and leaf water potential were both lower than at the coastal site. E. californica shrubs were leafless much of the time under natural conditions in the desert garden and had the smallest size there as well. Under natural conditions, E. farinosa, with its reflective leaf surface, was able to maintain lower leaf temperatures and attained a large size than the other two species in the desert garden. The green-leaved species (E. californica and E. frutescens) were not able to maintain leaves into the drought periods in the desert garden, with the exception of the irrigated E. frutescens which did maintain its leaf area if provided with supplemental watering to maintain transpirational leaf cooling. In the coastal garden, all species survived and there were few clear differences in the physiological characteristics among the three species. E. californica, the coastal native, attained a larger size in the coastal garden when compared with either of the two desert species.     

Development of leaf water deficits in detached green and lime-chlorotic leaves of seedlings from populations ofEucalyptus obliqua L'Hérit

Summary The water relations of three populations ofEucalyptus obliqua affected by limechlorosis were studied by monitoring water loss of leaves detached from seedlings. Water was lost faster from chlorotic leaves than from green leaves of the same population; the rate of water loss was inversely propotional to the chlorophyll content. Although chlorophyll content may affect stomatal behaviour, the water loss curves were interpreted as indicating that cuticular aspects of leaf water retention were severely affected.These findings concur with recent observations of severe water stress in chlorotic plants growing on calcareous soils in the Cape Otway region. Moreover, it appears that the greater susceptibility of the acidic population to lime-chlorosis compared with that of the calcareous population would be a considerable disadvantage when growing on a calcareous site subjected to periodic droughts.     

Leaf phenology, seasonal changes in leaf quality and herbivory pattern of Sanguisorba tenuifolia at different altitudes

Leaf demography, seasonal changes in leaf quality and leaf-beetle herbivory of a herbaceous perennial plant, Sanguisorba tenuifolia, were compared between low- and high-elevation sites. Leaf nitrogen concentration was higher and leaf mass per area (LMA) was lower at the higher site than at the lower one. At the lower site, with a long growth period, plants produced many leaf cohorts and leaves emerged throughout the growing season. At the higher site, with a short growth period, however, leaf emergence was concentrated early in the growing season. The improvement of leaf quality and acceleration of leaf emergence at higher altitude are seen as adaptations to a short growing season. Results of a feeding trial suggested that leaf quality for the leaf-beetle Galerucella grisescens was higher at the higher site, but plants at the higher site showed less damage. Oviposition of G. grisescens was seasonal and unimodal at both altitudes, but the period of oviposition was shorter and its density lower at the higher site. The low temperature and short growth period at the higher site appear to reduce the activity of the leaf-beetles, resulting in a decrease in damage by herbivory, despite better leaf quality. Received: 11 December 1997 / Accepted: 24 July 1998     

Leaf hairs: Effects on physiological activity and adaptive value to a desert shrub

Summary The effects of leaf hairs on photosynthesis, transpiration, and leaf energy balance were measured on the desert shrub Encelia farinosa in order to determine the adaptive significance of the hairs. The pubescence reduces leaf absorptance resulting in a reduced heat load, and as a consequence lower leaf temperatures and lower transpiration rates. In its native habitat where air temperatures often exceed 40° C, the optimum temperature for photosynthesis in E. farinosa occurs at 25° C, and at leaf temperatures above 35° C net photosynthesis declines precipitously. An advantage of leaf pubescence is that it allows a leaf temperature much lower than air temperature. As a result, leaf temperatures are near the temperature optimum for photosynthesis and high, potentially lethal leaf temperatures are avoided. However, there is a disadvantage associated with leaf pubescence. By reflecting quanta that might otherwise be used in photosynthesis, the presence of leaf hairs reduces the rate of photosynthesis. A tradeoff model was used to assess the overall advantage of possessing leaf hairs. In terms of the carbon gaining capacity of the leaf, the model predicted that for different environmental conditions different levels of leaf pubescence were optimal. In other words, under aird conditions and/or high air temperatures, leaves of E. farinosa would have a higher rate of photosynthesis by being pubescent than by not being pubescent. The predictions from this model agreed closely with observed patterns of leaf pubescence in the field.C.I.W.-D.P.B. Publication No. 613     

Comparative ecophysiology of Encelia farinosa and Encelia frutescens

Summary Encelia farinosa and Encelia frutescens are drought-decidous shrubs whose distributions overlap throughout much of the Sonoran Desert. During hot and dry periods, leaves of E. farinosa utilize increased leaf reflectance to reduce leaf temperature, whereas leaves of E. frutescens have substantially higher leaf conductances and rely on increased transpirational cooling to reduce leaf temperature. E. farinosa is common on the dry slope microhabitats, whereas E. frutescens occurs only in wash microhabitats where greater soil moisture is available to provide the water necessary for transpirational cooling. E. farinosa tends not to persist in wash microhabitats because of its greater susceptibility to flashfloods. The consequences and significance of increased leaf reflectance versus increased transpirational cooling to leaf temperature regulation are discussed.     

Combined effects of light and water availability on photosynthesis and growth of Arisaema heterophyllum in the forest understory and an open site

Photosynthetic characteristics, leaf longevity and biomass accumulation of a threatened herb species, Arisaema heterophyllum, were studied in the understory of a riparian forest and at a neighboring deforested open site for 3 years in order to understand the combined effects of light and water availability. Light availability was 2- to 4-fold higher at the deforested than at the forest site during the growing season of the species, and precipitation varied considerably over the 3 years. Despite the difference in water availability among the years (dry in 1994 and 1996, and wet in 1995), the species showed a strong acclimation to the different light environments. Light-saturated assimilation rate on a leaf area basis, leaf mass area ratio (LMA), and relative growth rate (RGR) were all higher at deforested site. While a positive correlation between individual RGR and microsite light availability was found in the wet year, no correlation was found in the dry years, and mean RGR was significantly lower in the dry year at both sites. Leaf longevity, photosynthetic capacity on a leaf mass basis, dark respiration rate, and leaf conductance, varied considerably from year to year, especially in the plants at the open site, probably depending on water availability. In the dry years plants at the deforested sites showed a lower photosynthetic rate and leaf conductance under unwatered than under watered conditions. These results suggest that the water availability in a given year may strongly affect light acclimation and annual RGR of the herb species in natural habitats, even under mesic climate conditions. Received: 15 February 1997 / Accepted: 20 May 1997     

Effects of growth temperature and winter duration on leaf phenology of Erythronium japonicum,a forest spring geophyte

The effects of growth temperature and winter duration on the leaf phenology of Erythronium japonicum were examined in two experiments. Bulbs wintered in the field were cultivated at 10 and 20° C and the bulbs were cultivated at 15° C after chilling treatment at 3° C for 60 and 120 days and without chilling in winter. The plants cultivated at 20° C showed significantly earlier leaf emergence, a more rapid rate of leaf extension and shorter leaf longevity than those cultivated at 10° C. The decrease in the leaf longevity at 20° C resulted from the decreases in the durations of all of the developmental, mature, and senescent phases. The bulbs without chilling treatment did not sprout leaves and those with chilling treatment sprouted leaves. The increase in the length of chilling treatment from 60 to 120 days affected leaf phenology in same manner as the increase in the growth temperature from 10 to 20° C.     

Coordination between leaf and stem traits related to leaf carbon gain and hydraulics across 32 drought-tolerant angiosperms

We examined 15 traits in leaves and stems related to leaf C economy and water use for 32 co-existing angiosperms at ridge sites with shallow soil in the Bonin Islands. Across species, stem density was positively correlated to leaf mass per area (LMA), leaf lifespan (LLS), and total phenolics and condensed tannins per unit leaf N (N-based), and negatively correlated to leaf osmotic potential and saturated water content in leaves. LMA and LLS were negatively correlated to photosynthetic parameters, such as area-, mass-, and N-based assimilation rates. Although stem density and leaf osmotic potential were not associated with photosynthetic parameters, they were associated with some parameters of the leaf C economy, such as LMA and LLS. In the principal component (PCA) analysis, the first three axes accounted for 74.4% of total variation. Axis 1, which explained 41.8% of the total variation, was well associated with parameters for leaf C and N economy. Similarly, axis 2, which explained 22.3% of the total variation, was associated with parameters for water use. Axis 3, which explained 10.3% of the total variation, was associated with chemical defense within leaves. Axes 1 and 2 separated functional types relatively well, i.e., creeping trees, ruderal trees, other woody plants, C₃ shrubs and forbs, palms, and CAM plants, indicating that plant functional types were characterized by similar attributes of traits related to leaf C and N economy and water use. In addition, when the plot was extended by two unrelated traits, leaf mass-based assimilation rates and stem density, it also separated these functional types. These data indicate that differences in the functional types with contrasting plant strategies can be attributed to functional integration among leaf C economy, hydraulics, and leaf longevity, and that both leaf mass-based assimilation rates and stem density are key factors reflecting the different functions of plant species.     

Best hyperspectral indices for tracing leaf water status as determined from leaf dehydration experiments

Leaf water status information is highly needed for monitoring plant physiological processes and assessing drought stress. Retrieval of leaf water status based on hyperspectral indices has been shown to be easy and rapid. However, a universal index that is applicable to various plants remains a considerable challenge, primarily due to the limited range of field-measured datasets. In this study, a leaf dehydration experiment was designed to obtain a relatively comprehensive dataset with ranges that are difficult to obtain in field measurements. The relative water content (RWC) and equivalent water thickness (EWT) were chosen as the surrogates of leaf water status. Moreover, five common types of hyperspectral indices including: single reflectance (R), wavelength difference (D), simple ratio (SR), normalized ratio (ND) and double difference (DDn) were applied to determine the best indices. The results indicate that values of original reflectance, reflectance difference and reflectance sensitivity increased significantly, particularly within the 350–700 nm and 1300–2500 nm domains, with a decrease in leaf water. The identified best indices for RWC and EWT, when all the species were considered together, were the first derivative reflectance based ND type index of dND (1415, 1530) and SR type index of dSR (1530, 1895), with R² values of 0.95 (p < 0.001) and 0.97 (p < 0.001), respectively, better than previously published indices. Even so, different best indices for different species were identified, most probably due to the differences in leaf anatomy and physiological processes during leaf dehydration. Although more plant species and field-measured datasets are still needed in future studies, the recommend indices based on derivative spectra provide a means to monitor drought-induced plant mortality in temperate climate regions.     

INFLUENCE OF IRRADIATION,SOIL WATER POTENTIAL,AND LEAF TEMPERATURE ON LEAF MORPHOLOGY OF A DESERT BROADLEAF,ENCELIA FARINOSA GRAY (COMPOSITAE)

Laboratory experiments were performed to evaluate observed seasonal changes in leaf morphology of the desert perennial shrub, Encelia farinosa Gray. Plants were grown under low or high conditions of photosynthetically active irradiation, soil water potential (Ψ^soil), and leaf temperature (8 different experimental regimes). The relative growth rate, leaf water vapor conductance, leaf water potential, and leaf length were all greater for the high Ψ^soil regimes, the largest leaves occurring at low irradiation. High irradiation during growth led to thicker leaves with a higher internal to external leaf area ratio (A^mes/A); low Ψ^soil tended to increase A^mes/A somewhat. High irradiation also led to decreased absorptance to solar irradiation caused by increased pubescence. High leaf temperature during development resulted in slightly smaller, thicker leaves with higher A^mes/A. Thus, irradiation appeared to have its major influence on leaf thickness, A^mes/A, and absorptance, with a secondary effect on leaf length; Ψ^soil affected primarily leaf length, growth rate, and water status, and secondarily A^mes/A. Results are discussed with regard to recent ecophysiological studies on the observed seasonal changes in leaf morphology of E. farinosa.     

Effects of growth temperature and winter duration on leaf phenology of a spring ephemeral (Gagea lutea) and a summergreen forb (Maianthemum dilatatum)

Effects of growth temperature and winter duration on leaf longevity were compared between a spring ephemeral, Gagea lutea, and a forest summergreen forb, Maianthemum dilatatum. The plants were grown at day/night temperatures of 25/20°C and 15/10°C after a chilling treatment for variable periods at 2°C. The temperature regime of 25/20°C was much higher than the mean air temperatures for both species in their native habitats. Warm temperature of 25/20°C and/or long chilling treatment shortened leaf longevity in G. lutea, but not in M. dilatatum. The response of G. lutea was consistent with that reported for other spring ephemerals. Air temperature increases as the vegetative season progresses. The decrease in leaf longevity in G. lutea under warm temperature condition ensures leaf senescence in summer, an unfavorable season for its growth. This also implies that early leaf senescence could occur in years with early summers. Warm spring temperatures have been shown to accelerate the leafing-out of forest trees. The decrease in leaf longevity due to warm temperature helps synchronize the period of leaf senescence roughly with the time of the forest canopy leaf-out. Prolonged winter due to late snowmelt has been shown to shorten the vegetative period for spring ephemerals. The decrease in leaf longevity due to long chilling treatment would correspond with this shortened vegetative period.     

Seasonal change of Lepidopteran larval assemblage associated with leaf quality

The community structure of herbivorous animals is frequently regulated by the nutritional condition of their host plant. The present study was aimed at investigating relationship between leaf chemistry traits and assemblages of Lepidoptera larvae in southwestern part of Korea. We selected three sites in rural landscapes and lepidopteran larvae were collected using a beating sheet from May to August. We collected leaves of two major host plants, Eurya japonicaThunb andQuercusserrataThunb and measured water content, nitrogen content, carbon content and area eaten of leaf. During the last four months we collected a total of 481 individuals in more than 25 different plant species. The number of larvae was highest in July and most abundant in Eurya japonica with 253 individuals. Leaf quality varied in each month. The correlation between leaf quality and assemblage of larvae showed that the carbon content only affected the number of larvae (r =?0.833, p < 0.01). Interestingly the water content and area eaten by the larvae was positively correlated (r = 0.835, p < 0.05).