1 Plant ecology at high elevations.- The concept of limitation.- A regional and historical account.- The challenge of alpine plant research.- 2 The alpine life zone.- Altitudinal boundaries.- Global alpine land area.- Alpine plant diversity.- Origin of alpine floras.- Alpine growth forms.- 3 Alpine climate.- Which alpine climate.- Common features of alpine climates.- Regional features of alpine climates.- 4 The climate plants experience.- Interactions of relief, wind and sun.- How alpine plants influence their climate.- The geographic variation of alpine climate.- 5 Life under snow: protection and limitation.- Temperatures under snow.- Solar radiation under snow.- Gas concentrations under snow.- Plant responses to snowpack.- 6 Alpine soils.- Physics of alpine soil formation.- The organic compound.- The interaction of organic and inorganic compounds.- 7 Alpine treelines.- About trees and lines.- Current altitudinal positions of climatic treelines.- Treeline-climate relationships.- Intrazonal variations and pantropical plateauing of alpine treelines.- Treelines in the past.- Attempts at a functional explanation of treelines.- A hypothesis for treeline formation.- Growth trends near treelines.- Evidence for sink limitation.- 8 Climatic stress.- Survival of low temperature extremes.- Avoidance and tolerance of low temperature extremes.- Heat stress in alpine plants.- Ultraviolet radiation - a stress factor.- 9 Water relations.- Ecosystem water balance.- Soil moisture at high altitudes.- Plant water relations - a brief review of principles.- Water relations of alpine plants.- Desiccation stress.- Water relations of special plant types.- 10 Mineral nutrition.- Soil nutrients.- The nutrient status of alpine plants.- Nutrient cycling and nutrient budgets.- Nitrogen fixation.- Mycorrhiza.- Responses of vegetation to variable nutrient supply.- 11 Uptake and loss of carbon.- Photosynthetic capacity of alpine plants.- Photosynthetic responses to the environment.- Daily carbon gain of leaves.- The seasonal carbon gain of leaves.- C4 and CAM photosynthesis at high altitudes.- Tissue respiration of alpine plants.- Ecosystem carbon balance.- 12 Carbon investments.- Non-structural carbohydrates.- Lipids and energy content.- Carbon costs of leaves and roots.- Whole plant carbon allocation.- 13 Growth dynamics and phenology.- Seasonal growth.- Diurnal leaf extension.- Rates of plant dry matter accumulation.- Functional duration of leaves and roots.- 14 Cell division and tissue formation.- Cell size and plant size.- Mitosis and the cell cycle.- From meristem activity to growth control.- 15 Plant biomass production.- The structure of alpine plant canopies.- Primary productivity of alpine vegetation.- Plant dry matter pools.- Biomass losses through herbivores.- 16 Plant reproduction.- Flowering and pollination.- Seed development and seed size.- Germination.- Alpine seed banks and natural recruitment.- Clonal propagation.- Alpine plant age.- Community processes.- 17 Global change at high elevation.- Alpine land use.- The impact of altered atmospheric chemistry.- Climatic change and alpine ecosystems.- References (with chapter annotation).- Taxonomic index (genera).- Geographical index.- Color plates.- Plant life forms.- The alpine life zone.- Environmental stress.- The human dimension.

Alpine plant life

Mangroves are woody plants that grow at the interface between land and sea in tropical and sub-tropical latitudes where they exist in conditions of high salinity, extreme tides, strong winds, high temperatures and muddy, anaerobic soils. There may be no other group of plants with such highly developed morphological and physiological adaptations to extreme conditions.

Because of their environment, mangroves are necessarily tolerant of high salt levels and have mechanisms to take up water despite strong osmotic potentials. Some also take up salts, but excrete them through specialized glands in the leaves. Others transfer salts into senescent leaves or store them in the bark or the wood. Still others simply become increasingly conservative in their water use as water salinity increases Morphological specializations include profuse lateral roots that anchor the trees in the loose sediments, exposed aerial roots for gas exchange and viviparous waterdispersed propagules.

Mangroves create unique ecological environments that host rich assemblages of species. The muddy or sandy sediments of the mangal are home to a variety of epibenthic, infaunal, and meiofaunal invertebrates Channels within the mangal support communities of phytoplankton, zooplankton and fish. The mangal may play a special role as nursery habitat for juveniles of fish whose adults occupy other habitats (e.g. coral reefs and seagrass beds).

Because they are surrounded by loose sediments, the submerged mangroves' roots, trunks and branches are islands of habitat that may attract rich epifaunal communities including bacteria, fungi, macroalgae and invertebrates. The aerial roots, trunks, leaves and branches host other groups of organisms. A number of crab species live among the roots, on the trunks or even forage in the canopy. Insects, reptiles, amphibians, birds and mammals thrive in the habitat and contribute to its unique character.

Living at the interface between land and sea, mangroves are well adapted to deal with natural stressors (e.g. temperature, salinity, anoxia, UV). However, because they live close to their tolerance limits, they may be particularly sensitive to disturbances like those created by human activities. Because of their proximity to population centers, mangals have historically been favored sites for sewage disposal. Industrial effluents have contributed to heavy metal contamination in the sediments. Oil from spills and from petroleum production has flowed into many mangals. These insults have had significant negative effects on the mangroves.

Habitat destruction through human encroachment has been the primary cause of mangrove loss. Diversion of freshwater for irrigation and land reclamation has destroyed extensive mangrove forests. In the past several decades, numerous tracts of mangrove have been converted for aquaculture, fundamentally altering the nature of the habitat. Measurements reveal alarming levels of mangrove destruction. Some estimates put global loss rates at one million ha y−1, with mangroves in some regions in danger of complete collapse. Heavy historical exploitation of mangroves has left many remaining habitats severely damaged.

These impacts are likely to continue, and worsen, as human populations expand further into the mangals. In regions where mangrove removal has produced significant environmental problems, efforts are underway to launch mangrove agroforestry and agriculture projects. Mangrove systems require intensive care to save threatened areas. So far, conservation and management efforts lag behind the destruction; there is still much to learn about proper management and sustainable harvesting of mangrove forests.

Mangroves have enormous ecological value. They protect and stabilize coastlines, enrich coastal waters, yield commercial forest products and support coastal fisheries. Mangrove forests are among the world's most productive ecosystems, producing organic carbon well in excess of the ecosystem requirements and contributing significantly to the global carbon cycle. Extracts from mangroves and mangrove-dependent species have proven activity against human, animal and plant pathogens. Mangroves may be further developed as sources of high-value commercial products and fishery resources and as sites for a burgeoning ecotourism industry. Their unique features also make them ideal sites for experimental studies of biodiversity and ecosystem function. Where degraded areas are being revegetated, continued monitoring and thorough assessment must be done to help understand the recovery process. This knowledge will help develop strategies to promote better rehabilitation of degraded mangrove habitats the world over and ensure that these unique ecosystems survive and flourish.

Biology of mangroves and mangrove Ecosystems

Stable isotopes in tree rings.

/pdf/hydrological-functions-of-tropical-forests-not-seeing-the-2hpleaj4on.pdf

Hydrological functions of tropical forests: not seeing the soil for the trees?

In this review I first compile data for the worldwide position of climate-driven alpine treelines. Causes for treeline formation are then discussed with a global perspective. Available evidence suggests a combination of a general thermal boundary for tree growth, with regionally variable “modulatory” forces, including the presence of certain taxa. Much of the explanatory evidence found in the literature relates to these modulatory aspects at regional scales, whereas no good explanations emerged for the more fundamental global pattern related to temperature per se, on which this review is focused. I hypothesize that the life form “tree” is limited at treeline altitudes by the potential investment, rather than production, of assimilates (growth as such, rather than photosynthesis or the carbon balance, being limited). In shoots coupled to a cold atmosphere, meristem activity is suggested to be limited for much of the time, especially at night. By reducing soil heat flux during the growing season the forest canopy negatively affects root zone temperature. The lower threshold temperature for tissue growth and development appears to be higher than 3°C and lower than 10°C, possibly in the 5.5–7.5°C range, most commonly associated with seasonal means of air temperature at treeline positions. The physiological and developmental mechanisms responsible have yet to be analyzed. Root zone temperature, though largely unknown, is likely to be most critical.

/pdf/a-re-assessment-of-high-elevation-treeline-positions-and-2h8ncswvxc.pdf

A re-assessment of high elevation treeline positions and their explanation

Andean cloud forests play an important role in watershed hydrology and protection against erosion. Even though most cloud forests fall under officially protected areas, a good deal of the cloud forests are being deforested and replaced by pastures for grazing cattle, which is the most important land use in Venezuelan cloud forest environments. The water dynamics of the natural forest as well as the impact of replacement by pastures are poorly understood. We have been conducting a research project since 1996, in order to study some of the water fluxes of the forest and to evaluate the hydrological impact of replacement by pasture. The study site is located at La Mucuy (2300 m with 3124 mm rainfall), Merida State, in the Venezuelan Andes. The results show that, in the forest, 91% of total incoming water was from rainfall and 9% was from cloudwater. Total foliage interception was 51%, which is a high value for a tropical montane forest. About 49% of the total incoming water reaches the ground as thr...

/pdf/deforestation-impact-on-water-dynamics-in-a-venezuelan-5837ee2iut.pdf

Deforestation Impact on Water Dynamics in a Venezuelan Andean Cloud Forest

Freezing tolerance and avoidance were studied in several different sized species of the tropical high Andes (4200 m) to determine whether there was a relationship between plant height and cold resistance mechanisms. Freezing injury and supercooling capacity were determined in ground level plants (i.e. cushions, small rosettes and a perennial herb), intermediate height plants (shrubs and perennial herbs) and arborescent forms (i.e. giant rosettes and small trees). All ground-level plants showed tolerance as the main mechanism of resistance to cold temperatures. Arborescent forms showed avoidance mechanisms mainly through supercooling, while intermediate plants exhibited both. Insulation mechanisms to avoid low temperatures were present in the two extreme life-forms. We suggest that a combination of freezing tolerance and avoidance by insulation is least expensive and is a more secure mechanism for high tropical mountain plants than supercooling alone.

Freezing tolerance and avoidance in high tropical Andean plants: Is it equally represented in species with different plant height?

Factors affecting supercooling capacity and cold hardiness were investigated in leaves of ten giant rosette species of the genus Espeletia (Compositae). These species grow along a 2,800-4,200 m elevation gradient in the Venezuelan Andes. In this high tropical environment, freezing frequently occurs every night, particularly above 3,300 m, but lasts for only a few hours. Supercooling capacty is linearly related to leaf water potential (Ψ L ) in all species; however supercooling is more responsive to Ψ L changes in Espeletia species from high paramos. The rate of change in the species-specific supercooling point and the rate of change of average annual minimum temperature along the elevation and climatic gradient follow the same trend (approximately -0.6 K per 100 m elevation). At a given elevation, the expanded leaves of the different species tend to supercool 8-10 K below minimum air temperatures. Experimentally-induced freezing was accompanied by the formation of intracellular ice and tissue damage. The relative apoplastic water content (RAWC) of the leaves, which may influence the ice nucleation rate or the facility by which ice propagates, was determined by pressure-volume methods. Species from higher sites tend to exhibit lower RAWC (2%-7%) than species from lower sites (7%-36%). A causal relationship between supercooling capacity and RAWC is suggested. In the high tropical Andes, the temperature oxotherm plateau of Espeletia leaves seems to be sufficiently fow to avoid freezing.

Cold hardiness and supercooling along an altitudinal gradient in andean giant rosette species.

The Polylepis tarapacana forests found in Bolivia are unique with respect to their altitudinal distribution (4200–5200 m). Given the extreme environmental conditions that characterize these altitudes, this species has to rely on distinct mechanisms to survive stressful temperatures. The purpose of this study was to determine low-temperature resistance mechanisms in P. tarapacana. Tissue was sampled for carbohydrate and proline contents and micro-climatic measurements were made at two altitudes, 4300 and 4850 m, during both the dry cold and wet warm seasons. Supercooling capacity (−3 to −6 °C for the cold dry and −7 to −9 °C for the wet warm season) and injury temperatures (−18 to −23 °C for both seasons), determined in the laboratory, indicate that P. tarapacana is a frost-tolerant species. On the other hand, an increase in supercooling capacity, as the result of significant increase in total soluble sugar and proline contents, occurs during the wet warm season as a consequence of higher metabolic activity. Hence, P. tarapacana, a frost-tolerant species during the colder unfavourable season, is able to avoid freezing during the more favourable season when minimum night-time temperatures are not as extreme.

Low‐temperature resistance in Polylepis tarapacana, a tree growing at the highest altitudes in the world

The effects of temperature on photosynthesis of a rosette plant growing at ground level, Acaena cylindrostachya R. et P., and an herb that grows 20–50 cm above ground level, Senecio formosus H.B.K., were studied along an altitudinal gradient in the Venezuelan Andes. These species were chosen in order to determine – in the field and in the laboratory – how differences in leaf temperature, determined by plant form and microenvironmental conditions, affect their photosynthetic capacity. CO2 assimilation rates (A) for both species decreased with increasing altitude. For Acaena leaves at 2900 m, A reached maximum values above 9 μmol m−2 s−1, nearly twice as high as maximum A found at 3550 m (5.2) or at 4200 m (3.9). For Senecio leaves, maximum rates of CO2 uptake were 7.5, 5.8 and 3.6 μmol m−2 s−1 for plants at 2900, 3550 and 4200 m, respectively. Net photosynthesis-leaf temperature relations showed differences in optimum temperature for photosynthesis (A
o.t.) for both species along the altitudinal gradient. Acaena showed similar A
o.t. for the two lower altitudes, with 19.1°C at 2900 m and 19.6°C at 3550 m, while it increased to 21.7°C at 4200 m. Maximum A for this species at each altitude was similar, between 5.5 and 6.0 μmol m−2 s−1. For the taller Senecio, A
o.t. was more closely related to air temperatures and decreased from 21.7°C at 2900 m, to 19.7°C at 3550 m and 15.5°C at 4200 m. In this species, maximum A was lower with increasing altitude (from 6.0 at 2900 m to 3.5 μmol m−2 s−1 at 4200 m). High temperature compensation points for Acaena were similar at the three altitudes, c. 35°C, but varied in Senecio from 37°C at 2900 m, to 39°C at 3550 m and 28°C at 4200 m. Our results show how photosynthetic characteristics change along the altitudinal gradient for two morphologically contrasting species influenced by soil or air temperatures.

Fermín Rada

Papers

Deforestation Impact on Water Dynamics in a Venezuelan Andean Cloud Forest

Freezing tolerance and avoidance in high tropical Andean plants: Is it equally represented in species with different plant height?

Cold hardiness and supercooling along an altitudinal gradient in andean giant rosette species.

Low‐temperature resistance in Polylepis tarapacana, a tree growing at the highest altitudes in the world

Effects of temperature on photosynthesis of two morphologically contrasting plant species along an altitudinal gradient in the tropical high Andes.