(Fig. 9) Pollen analysis of sediment core GIK16017-2

The improved understanding of the pollen signal in the marine sediments offshore of northwest Africa is applied to deep-sea core M 16017-2 at 21°N. Downcore fluctuations in the percentage, concentration and influx diagrams record latitudinal shifts of the main northwest African vegetation zones and characteristics of the trade winds and the African Easterly Jet. Time control is provided by 14C ages and 180 records. During the period 19,000-14,000 yr B.P. a compressed savanna belt extended between about 12 ° and 14-15°N. The Sahara had maximally expanded northward and southward under hyperarid climatic conditions. The belt with trade winds and dominant African Easterly Jet transport had not shifted latitudinally. The trade winds were strong as compared to the modern situation but around 13,000 yr B.P. the trade winds weakened. After 14,000 yr B.P. the climate became less arid south of the Sahara and a first spike of fluvial runoff is registered around 13,000 yr B.P. Fluvial runoff increased strongly around 11,000 yr B.P. and maximum runoff is recorded from about 9000-7800 yr B.P. Around 12,500 yr B.P. the savanna belt started to shift northward and became richer in woody species: it shifted about 6° of latitude, reached its northernmost position during the period of 9200-7800 yr B.P. and extended between about 16° and 24°N at that time. Tropical forest had reached its maximum expansion and the Guinea zone reached as far north as about 15°N, reflecting very humid climatic conditions south of the Sahara. North of the Sahara the climate also became more humid and Mediterranean vegetation developed rapidly. The Sahara had maximally contracted and the trade winds were weak and comparable with the present day intensity. After about 7800 yr B.P. the southern fringe of the Sahara and accordingly the savanna belt, shifted rapidly southward again.

Adventitious root formation in cuttings of Backhousia citriodora F. Muell: 1. Plant genotype, juvenility and characteristics of cuttings

Backhousia citriodora is a commercially valuable Australian woody species that has a reputation for being recalcitrant in forming adventitious roots from cuttings. A study was carried out to determine whether maturation and plant genotype influenced rooting. It also tried to establish whether genotypic differences in rooting ability were related to characteristics of the cutting material. The rooting of cuttings in B. citriodora declines after maturation and is strongly influenced by genotype. The cutting characteristics of actively growing axillary buds, wide stems and mature leaves are associated with rooting and survival but not related to genotype. Furthermore, the 8-24 weeks required by B. citriodora to form roots from cuttings makes it difficult to distinguish between the characteristics that increase rooting and those characteristics that enhance survival. A subsequent disbudding experiment demonstrated that axillary buds per se have an inhibitory effect on rooting. This suggests that the presence of actively growing axillary buds are an indication of overall growth and condition of the stock plant unrelated to the formation of adventitious rooting. The effects of other cutting characteristics on rooting are also discussed. (C) 2004 Elsevier B.V. All rights reserved.

Why Leaves Turn Red in Autumn. The Role of Anthocyanins in Senescing Leaves of Red-Osier Dogwood

Why the leaves of many woody species accumulate anthocyanins prior to being shed has long puzzled biologists because it is unclear what effects anthocyanins may have on leaf function. Here, we provide evidence for red-osier dogwood (Cornus stolonifera) that anthocyanins form a pigment layer in the palisade mesophyll layer that decreases light capture by chloroplasts. Measurements of leaf absorbance demonstrated that red-senescing leaves absorbed more light of blue-green to orange wavelengths (495–644 nm) compared with yellow-senescing leaves. Using chlorophyll a fluorescence measurements, we observed that maximum photosystem II (PSII) photon yield of red-senescing leaves recovered from a high-light stress treatment, whereas yellow-senescing leaves failed to recover after 6 h of dark adaptation, which suggests photo-oxidative damage. Because no differences were observed in light response curves of effective PSII photon yield for red- and yellow-senescing leaves, differences between red- and yellow-senescing cannot be explained by differences in the capacities for photochemical and non-photochemical light energy dissipation. A role of anthocyanins as screening pigments was explored further by measuring the responses PSII photon yield to blue light, which is preferentially absorbed by anthocyanins, versus red light, which is poorly absorbed. We found that dark-adapted PSII photon yield of red-senescing leaves recovered rapidly following illumination with blue light. However, red light induced a similar, prolonged decrease in PSII photon yield in both red- and yellow-senescing leaves. We suggest that optical masking of chlorophyll by anthocyanins reduces risk of photo-oxidative damage to leaf cells as they senesce, which otherwise may lower the efficiency of nutrient retrieval from senescing autumn leaves.

Pigment dynamics and autumn leaf senescence in a New England deciduous forest, eastern USA

The leaves of woody plants at Harvard Forest in Central Massachusetts, USA, changed color during senescence; 70% (62/89) of the woody species examined anatomically contained anthocyanins during senescence. Anthocyanins were not present in summer green leaves, and appeared primarily in the vacuoles of palisade parenchyma cells. Yellow coloration was a result of the unmasking of xanthophyll pigments in senescing chloroplasts. In nine red-senescing species, anthocyanins were not detectable in mature leaves, and were synthesized de novo in senescence, with less than 20 m g cm - 2 of chlorophyll remaining. Xanthophyll concentrations declined in relation to chlorophyll to the same extent in both yellow- and red-leaved taxa. Declines in the maximum photosystem II quantum yield of leaves collected prior to dawn were only slightly less in the red-senescing species, indicating no long-term protective activity. Red-leaved species had significantly greater mass/area and lower chlorophyll a / b ratios during senescence. Nitrogen tissue concentrations in mature and senescent leaves negatively correlated to anthocyanin concentrations in senescent leaves, weak evidence for more efficient nitrogen resorption in anthocyanic species. Shading retarded both chlorophyll loss and anthocyanin production in Cornus alternifolia , Acer rubrum , Acer saccharum , Quercus rubra and Viburnum alnifolium . It promoted chlorophyll loss in yellow-senescing Fagus grandifolia . A reduced red : far-red ratio did not affect this process. Anthocyanins did not increase leaf temperatures in Q. rubra and Vaccinium corymbosum on cold and sunny days. The timing of leaf-fall was remarkably constant from year to year, and the order of senescence of individual species was consistent.

Vegetation dynamics and their implications for the management of wetlands in the lowlands of Nepal

The maintenance of species richness is often a priority in the management of nature reserves, where consumptive use of resources is generally prohibited. The purpose of this research was to improve management by understanding the vegetation dynamics in the lowlands of Nepal. The objectives were to determine vegetation associations in relation to environments and human-induced disturbances that affect vegetation dynamics on floodplains, where upstream barrages had altered flooding patterns, and consumptive use of plant resources was influencing natural processes. Floodplain vegetation in relation to physical environments and disturbances were studied along transects, perpendicular to the course of the Mahakali River in the western Terai, Nepal. Forest structural changes were studied for three years in ten plots. A randomized split-block experiment with nine burning and grazing treatments was performed in seasonally flooded grasslands. A semi-structured questionnaire was used to assess people's socio-economic status, natural resource use patterns and conservation attitudes. ^ Elevation, soil organic matter, nitrogen, percentage of sand and grazing intensity were significant in delineating herbaceous vegetation assemblages, whereas elevation and livestock grazing were significant in defining forest type boundaries. On the floodplain islands, highly grazed Dalbergia sissoo-Acacia catechu forests were devoid of understory woody vegetation, but the lightly grazed D. sissoo-mixed forests had a well-developed second canopy layer, comprising woody species other than D. sissoo and A. catechu. In grasslands, species richness and biomass production were highest at intermediate disturbance level represented by the lightly grazed and ungrazed early-burned treatments. Ethnicity, education and resource use patterns were important in influencing conservation attitudes. A succession towards the mixed forests would occur in D. sissoo-dominated floodplain forests, where dams and barrages reduce flooding and associated fluvial processes, and if livestock grazing is stopped, as occasionally suggested by nature conservationists. In seasonally flooded grasslands, early burning with moderate grazing would enhance the species diversity and productivity. There is a need to implement a participatory integrated wetland management plan, to include community development, education and off farm income generation, to assure participatory conservation and management of wetlands in Nepal. ^

Chilling damage in a changing climate in coastal landscapes of the subtropical zone: a case study from south Florida

Extensive portions of the southern Everglades are characterized by series of elongated, raised peat ridges and tree islands oriented parallel to the predominant flow direction, separated by intervening sloughs. Tall herbs or woody species are associated with higher elevations and shorter emergent or floating species are associated with lower elevations. The organic soils in this “Ridge-and-Slough” landscape have been stable over millennia in many locations, but degrade over decades under altered hydrologic conditions. We examined soil, pore water, and leaf phosphorus (P) and nitrogen (N) distributions in six Ridge and Slough communities in Shark Slough, Everglades National Park. We found P enrichment to increase and N to decrease monotonically along a gradient from the most persistently flooded sloughs to rarely flooded ridge environments, with the most dramatic change associated with the transition from marsh to forest. Leaf N:P ratios indicated that the marsh communities were strongly P-limited, while data from several forest types suggested either N-limitation or co-limitation by N and P. Ground water stage in forests exhibited a daytime decrease and partial nighttime recovery during periods of surface exposure. The recovery phase suggested re-supply from adjacent flooded marshes or the underlying aquifer, and a strong hydrologic connection between ridge and slough. We therefore developed a simple steady-state model to explore a mechanism by which a phosphorus conveyor belt driven by both evapotranspiration and the regional flow gradient can contribute to the characteristic Ridge and Slough pattern. The model demonstrated that evapotranspiration sinks at higher elevations can draw in low concentration marsh waters, raising local soil and water P concentrations. Focusing of flow and nutrients at the evapotranspiration zone is not strong enough to overcome the regional gradient entirely, allowing the nutrient to spread downstream and creating an elongated concentration plume in the direction of flow. Our analyses suggest that autogenic processes involving the effects of initially small differences in topography, via their interactions with hydrology and nutrient availability, can produce persistent physiographic patterns in the organic sediments of the Everglades.

Moringa oleifera plasticity to face climatic changes

Extreme abiotic factors, such drought combined with heat waves and/or high UVB radiation are predicted to become more frequent in the future. The impact on plant production of these challenges on multipurpose Moringa oleifera L. remains unclear. A susceptibility of this species may lead to increase poverty in endangered regions. M. oleifera is a woody species native from sub-Himalaya regions under high climate stress pressure. The interest on this species is emerging due to its several medicinal properties and its nutritional value. Agropharmaceutical industry is interest in this species too. To understand the impact of increased climate factors, young (2 months old) plants of this species were exposed to water deficit (WD) and UVB (alone or combined). WD and WD+UVB imposition consists of unwater for 4 days. After 1 day withholding water, UVB and WD+UVB were irradiated with 26.3 kJ m-2 distributed per 3 days. Immediately after treatment exposition (1 day) and after 10 days, plant water status, growth, carbon metabolism and oxidative stress were measured. Overall no significant differences were observed in WD, regarding the parameters analysed, except on gas exchanges, MDA and phenols. The plants exposed to UVB showed, in general, more severe effects, as higher pigment content, MDA and membrane permeability, while no changes were observed in the total antioxidant activity. Plants exposed to UVB+WD, despite changes observed, the impact was lower than the one observed in UVB exposed plants, meaning that a protective/adaptive mechanism was developed in the plants under combined stressors. On the other hand, in all treatments the net CO2 assimilation rate decreased. Results suggest that M. oleifera has some tolerance to WD and UVB, and that develops mechanism of adaptation to these two types of stress that often arise in combination under a climate change scenario.

Dinâmica da regeneração natural de áreas em restauração pela transposição de solo superficial de cerrado e de floresta estacional

Dissertação (mestrado)—Universidade de Brasília, Instituto de Ciências Biológicas, Departamento de Ecologia, Programa de Pós-Graduação em Ecologia, 2015.

Pesticide bioconcentration modelling for fruit trees.

The model presented allows simulating the pesticide concentration in fruit trees and estimating the pesticide bioconcentration factor in fruits of woody species. The model allows estimating the pesticide uptake by plants through the water transpiration stream and also the time in which maximum pesticide concentration occur in the fruits. The equation proposed presents the relationships between bioconcentration factor (BCF) and the following variables: plant water transpiration volume (Q), pesticide transpiration stream concentration factor (TSCF), pesticide stem-water partition coefficient (KWood,w), stem dry biomass (M) and pesticide dissipation rate in the soil-plant system (kEGS). The modeling started and was developed from a previous model ?Fruit Tree Model? (FTM), reported by Trapp and collaborators in 2003, to which was added the hypothesis that the pesticide degradation in the soil follows a first order kinetic equation. The model fitness was evaluated through the sensitivity analysis of the pesticide BCF values in fruits with respect to the model entry data variability.

Assessing canopy temperature patterns in two grapevine varieties subjected to deficit irrigation: a tool to optimize water management ?

A better understanding of grapevine responses to drought and high air temperatures can help to optimize vineyard management to improve water use efficiency, yield and berry quality. Faster and robust field phenotyping tools are needed in modern precision viticulture, in particular in dry and hot regions such as the Mediterranean. Canopy temperature (Tc) is commonly used to monitor water stress in plants/crops and to characterize stomatal physiology in different woody species including Vitis vinifera. Thermography permits remote determination of leaf surface or canopy temperature in the field and also to assess the range and spatial distribution of temperature from different parts of the canopies. Our hypothesis is that grapevine genotypes may show different Tc patterns along the day due to different stomatal behaviour and heat dissipation strategies. We have monitored the diurnal and seasonal course of Tc in two grapevine genotypes, Aragonez (syn. Tempranillo) and Touriga Nacional subjected to deficit irrigation under typical Mediterranean climate conditions. Temperature measurements were complemented by determination of the diurnal course of leaf water potential (ψleaf) and leaf gas exchange. Measurements were done in two seasons (2013 and 2014) at different phenological stages: i) mid-June (green berry stage), ii) mid-July (veraison), iii) early August (early ripening) and iv) before harvest (late ripening). Correlations between Tc and minimal stomatal conductance will be presented for the two genotypes along the day. Results are discussed over the use of thermal imagery to derive information on genotype physiology in response to changing environmental conditions and to mild water stress induced by deficit irrigation. Strategies to optimize the use of thermal imaging in field conditions are also proposed

Differences between root traits of early- and late-successional trees influence below-ground competition and seedling establishment.

The competitive influence of the root system of the exotic grass Urochloa brizantha and the widespread forb Leonotis nepetifolia on the emergence, survival and early growth of the seedlings of eight tropical heliophilous herbaceous species, six early-successional woody species and five late-successional woody species from Brazil, grown in 3500-cm3 pots and in greenhouse without light restriction were assessed. The density of fine-root systems produced by the forb and the grass in pots were 6.8 cm cm-3 soil and 48.1 cm cm-3 soil, respectively. Seedlings survival of the heliophilous herbaceous, early- and late-successional woody species were 86%, 70% and 100% in presence of the forb root system and 12%, 14% and 100% in competition with grass root system, respectively. The competitive pressure applied by the grass root system on seedling growth of the heliophilous herbaceous, early- and late-successional woody species were 2.4, 1.9 and 1.4 times greater than the forb root system. Total root length of the heliophilous herbaceous, early- and late-successional woody species grown without competitors were 13, 33 and 5 times greater than in competition with forb, and were 66, 54 and 6 times greater than in competition with grass root system, respectively. The averages of fine-root diameter of plants grown without competitors were 209 microm for the heliophilous herbaceous, 281 microm for early-successional trees and 382 microm for late-successional trees. The root system of the forb did not avoid seedling establishment of most plant species, but the grass root system hampered more the establishment of heliophilous herbaceous and early-successional woody species than the seedling establishment of late-successional woody species. The different density of root systems produced in soil by the forb and the grass, and the distinct root traits (e.g. root diameter and root tissue density) of the early- and late-successional plant species can explain the differences in the establishment of seedlings of plant species belonging to different groups of tropical succession when exposed to below-ground competition.

Produção e degradação da serrapilheira de espécies lenhosas da caatinga e sua contribuição na recuperação de áreas degradadas.

Resumo: O entendimento do fluxo de produção e do aporte de nutrientes via decomposição da serrapilheira e as interações do processo com parâmetros edáficos e ciclagem de nutrientes de espécies nativas da Caatinga têm sido pouco estudados. O conhecimento sobre ciclagem de nutrientes em florestas manejadas também permite inferências sobre as espécies com maior capacidade de reciclagem de nutrientes e seu potencial para recuperação de áreas degradadas. Objetivou-se com isso avaliar a produção e a degradação da serrapilheira de oito espécies lenhosas da Caatinga e mensurar os efeitos de sua aplicação sobre a fertilidade do solo e sobre a produção de sorgo em solo degradado. Para isso realizou-se três ensaios: para o ensaio I quantificou-se a produção de serrapilheira em um delineamento inteiramente casualizado com 6 repetições, por meio da instalação de coletores sob a projeção da copa das espécies (tratamentos): mofumbo, sabiá, jurema-preta, jucá, catingueira, pereiro, pau-branco e marmeleiro, sendo o material coletado mensalmente; foram quantificadas a produção das frações folhas, caule, material reprodutivo, miscelânea e total, bem como o aporte de nutrientes no período chuvoso e seco. Para o ensaio II avaliou-se a taxa de degradação da fração folhas de cada espécie citada por meio da utilização de litter bags, em delineamento inteiramente casualizado com 4 repetições, as coletas foram aos 0, 30, 60, 90, 120 e 150 dias, em seguida quantificou-se os macro e micronutrientes, celulose, lignina e carbono em cada tempo de amostragem. Para o ensaio III, realizou-se experimento em casa de vegetação para mensurar os efeitos da aplicação dos resíduos da serrapilheira das mesmas espécies mencionadas nos ensaios anteriores (I e II) sobre a fertilidade do solo e a produção de sorgo em solo degradado, neste experimento adotou-se o delineamento em blocos casualizados com 5 tratamentos e 5 repetições, sendo avaliadas doses equivalentes a: 0, 15, 30, 60 e 120 kg ha-1 de N dos resíduos de cada espécie e um tratamento adicional com adubação mineral, totalizando 30 unidades experimentais para cada espécie. As variáveis mensuradas foram biométricas, biomassa, teor relativo de clorofila e nitrogênio total, além de análises de fertilidade do solo. Com a análise dos dados verificou-se que a época de maior produção de serrapilheira ocorreu no final do período chuvoso para o início do período seco. A espécie jucá apresentou maior produção de serrapilheira, comparado às outras espécies. O nutriente cálcio apresentou maior acúmulo na serrapilheira para as espécies mofumbo, sabiá, catingueira, pereiro e marmeleiro e o nitrogênio foi superior para as espécies jurema-preta, jucá e pau-branco. Para todas as espécies avaliadas no ensaio de degradação houve redução significativa na sua biomassa em relação ao tempo zero, apresentando a seguinte ordem de velocidade de decomposição: jurema-preta > catingueira > pau-branco > jucá > marmeleiro > mofumbo > pereiro > sabiá. No ensaio de fertilização com os resíduos verificou-se que o marmeleiro promoveu efeitos negativos no solo, como acidificação. Porém, a aplicação dos resíduos da espécie pau-branco foi a que promoveu aumento nos valores de K, SB e CEC do solo e na produção do sorgo os resíduos de jurema-preta e pau-branco foram as que promoveram aumento na massa seca das plantas. Enquanto a adubação mineral proporcionou aumento na produção de massa seca do sorgo, demonstrando que a associação entre adubo mineral e o uso da serrapilheira de espécies da Caatinga pode ser uma opção viável para acelerar a recuperação de solos degradados. Abstract: The understanding of the production flow and nutrient supply via decomposition of litter and process interactions with edaphic parameters and nutrient cycling of native species of the Caatinga has been little studied. The knowledge of nutrient cycling in managed forests also allow inferences about species with capacity greater nutrient recycling capacity and its potential for recovery of degraded areas. This study aimed to evaluate the production and litter degradation 8 woody species of Caatinga and measure the effects of its application on soil fertility and production of sorghum in degraded soil. To this was carried out three tests: for the test I quantified the production of litter in a completely randomized design with 6 replications, by installing collectors under the canopy projection in the species (treatments): mofumbo, sabiá, jurema-preta, jucá, catingueira, pereiro, pau-branco and marmeleiro for each species, and the material collected monthly, were quantified the production of fractions leaves, stem, reproductive material, miscellany and total nutrient intake in the rainy and dry season. For II test evaluated the degradation rate of the fraction leaves through the use of litter bags, in a completely randomized design with 4 replications, the collected was 0, 30, 60, 90, 120 and 150 days and quantitated nutrients, cellulose, lignin and carbon at each evaluation time. For the III test, there was the experiment in a greenhouse to measure the effects of the application of litter waste of the same species of previous tests (I and II) on soil fertility and production of sorghum in degraded soil, was adopted the randomized block design with 5 treatments and 5 replications and evaluated doses equivalent to: 0, 15, 30, 60 and 120 kg ha-1 N of waste each species and an additional treatment with mineral fertilizer, totaling 30 experimental units for each species. Biometric analysis and biomass, relative chlorophyll content and total nitrogen were proceeded. In addition to soil fertility analysis. With the data analysis it was found that the time of greatest litterfall occurred at the end of the rainy season to the beginning of the dry season. The jucá species showed higher production compared to other species. The nutrient calcium had higher accumulation for the species mofumbo, sabiá, catingueira, pereiro and marmeleiro and nitrogen was higher for species jurema-preta, jucá and pau-branco. All species evaluated in degradation test had a significant reduction in biomass over time zero. They presented the following order of decomposition rate: jurema-preta > catingueira > pau-branco > jucá > marmeleiro > mofumbo > pereiro > sabiá. For fertility test it was found that marmeleiro promoted negative effects on soil, such as acidification. However, pau-branco was the specie that promoted further improvements in the K values, SB and CEC to the soil and for the production of sorghum, the waste jurema-preta and pau-branco promoted increase in dry matter plants. While the mineral fertilization provided an increase in dry matter production of sorghum, demonstrating that the combination of mineral fertilizer and the use of litter of Caatinga species may be a viable option to speed up the recovery of degraded soils.

Leaf trait co-ordination in relation to construction cost, carbon gain and resource-use efficiency in exotic invasive and native woody vine species

Background and Aims: Success of invasive plant species is thought to be linked with their higher leaf carbon fixation strategy, enabling them to capture and utilize resources better than native species, and thus pre-empt and maintain space. However, these traits are not well-defined for invasive woody vines. Methods: In a glass house setting, experiments were conducted to examine how leaf carbon gain strategies differ between non-indigenous invasive and native woody vines of south-eastern Australia, by investigating their biomass gain, leaf structural, nutrient and physiological traits under changing light and moisture regimes. Key Results: Leaf construction cost (CC), calorific value and carbon : nitrogen (C : N) ratio were lower in the invasive group, while ash content, N, maximum photosynthesis, light-use efficiency, photosynthetic energyuse efficiency (PEUE) and specific leaf area (SLA) were higher in this group relative to the native group. Trait plasticity, relative growth rate (RGR), photosynthetic nitrogen-use efficiency and water-use efficiency did not differ significantly between the groups. However, across light resource, regression analyses indicated that at a common (same) leaf CC and PEUE, a higher biomass RGR resulted for the invasive group; also at a common SLA, a lower CC but higher N resulted for the invasive group. Overall, trait co-ordination (using pair-wise correlation analyses) was better in the invasive group. Ordination using 16 leaf traits indicated that the major axis of invasive-native dichotomy is primarily driven by SLA and CC (including its components and/or derivative of PEUE) and was significantly linked with RGR. Conclusions: These results demonstrated that while not all measures of leaf resource traits may differ between the two groups, the higher level of trait correlation and higher revenue returned (RGR) per unit of major resource need (CC) and use (PEUE) in the invasive group is in line with their rapid spread where introduced.

The influence of powdery mildew infection on photosynthesis, chlorophyll fluorescence, leaf chlorophyll and carotenoid content of three woody plant species

The effect of powdery mildew development on photosynthesis, chlorophyll fluorescence, leaf chlorophyll and carotenoid concentrations on three woody plants frequently planted in urban environments was studied. Rates of photosynthetic CO2 fixation were rapidly reduced in two of the three genotypes tested prior to visible signs of infection. Effects on chlorophyll fluorescence (Fo, Fv/Fo, Fv/Fm), leaf chlorophyll and carotenoid content were not manifest until >25 per cent of the leaf area was observed to be covered by mycelial growth indicating reduced photo-synthetic rates during the early stages of infection were not due to degradation of the leaf chloroplast structure. Observation of the fluorescence transient (OJIP curves) showed powdery mildew infection impairs photosynthetic electron transport system by reducing the size but not heterogeneity of the plastoquninone pool, effecting both the acceptor and donor side of photosystem II. Impairment of the photosynthetic electron transport system was reflected by reduced values of a performance index used in this investigation as a measure of photochemical events within photosystem II electron transport. In addition interpretation of the fluorescence data indicated powdery mildew infection may impair the photo-protective process that facilitates the dissipation of excess energy within leaf tissue.