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Alpine Plant Life: Functional Plant Ecology of High Mountain Ecosystems
Generations of plant scientists have been fascinated by alpine plant life - with the exposure of organisms to dramatic climatic gradients over a very short distance. This comprehensive text treats a wide range of alpine climate and soils, plant distribution and the treeline phenomenon, physiological ecology of water-, nutritional- and carbon relations of alpine plants, plant stress and plant development, biomass production, and aspects of human impacts on alpine vegetation. Geographically the book covers all parts of the world including the tropics.This second edition of Alpine Plant Life gives new references, new diagrams, and extensively revised chapters.
360 pages, Hardcover
First published January 1, 1999
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October 28, 2018
For those interested in alpine plants and their ecology, this is a great book on the physiology of the plants. The use of botanical terms is light, making the book very readable. Some highlights:
Chapter 1 - Plant ecology at high elevations
- conditions in the alpine provide natural experiments in comparative ecological research
Chapter 2 - The alpine life zone
- higher alpine plants grow up to 6000 m. in the Andes and Himalayas
- total alpine flora of the world is likely 8,000 to 10,000 higher plants
- alpine plants are typically prostrate woody shrubs, graminoids, cushion plants and herbaceous perennials often forming rosettes
- annuals commonly do not represent more than 2 percent of the alpine flora
Chapter 3 - Alpine climate
- while the lapse rate is usually taken to be 1 degree C per 100m, 0.6 degrees seems to be more typical for temperate summers and the tropics
- alpine plants can be subject to high radiation, both direct and diffuse from neighbouring clouds
Chapter 4 - The climate plants experience
- while conditions can be harsh in the alpine, it is different in the plant leaf canopy close to the ground where conditions may be 27 degrees C and 98 percent RH
- snow disappearance can vary with wind and exposure, varying by as much as 4 months - some plants can miss out one or two complete summers due to snow pack
- rosette plants can concentrate solar radiation, producing leaf temperatures over 50 degrees C
- Silene acaulis can produce leaf temperatures over 10 C degrees above air temperature; cushion plants become more abundant in windy environments
- similarly, tussock grasses use dead but sanding leaves to minimize wind effects
- root temperatures are low, often being only a few degrees above freezing
Chapter 5 - Life under snow
- alpines are able to photosynthesize at temperatures near freezing; some plants can reach 20 - 30 percent of their maximal rates
- as snow if not entirely reflective, plants are able to achieve net photosynthetic carbon gains under shallow snow
- even at 2 m. depth, photo signals are strong enough to induce seed germination, and sprouting
- CO2 concentrations under snow can exceed 1000 ppm, but this is in line with soil concentrations and do not appear to provide a significant benefit
- while oxygen concentration in snowpacks may be reduced, it does not appear to interfere with plant respiration
- the red colour of early leaves and stems is due to a lack of green chloroplasts allowing them to establish a radiation protective screen before the photosynthetic machinery is installed
- faster developing species prefer the center of snowbeds, while those taking more time prefer the edges
- it seems unlikely that sub-snow photosynthesis contributes significantly to the annual carbon balance of higher plants exposed to long-lasting snowpacks
Chapter 6 - Alpine soils
- dust accumulation has been found to have formed 10 cm of fine soil over the post-glacial period
- freezing is important in the heaving of bulk soils, resulting in mixing
- freeze-thaw causes downslope movement or solifluction, causing mechanical stress on the root systems
- accumulation of dead plant material results in biologically driven soil formation; essentially private recycling by cushion plants
- decomposition of leaf remains may take from 2 years for forbs to 5 years for sedges
- in the Rocky Mountains, it takes about 10,000 years for alpine soils to reach 30 cm
- the diversity of soil fungi decreases with altitude
Chapter 7 - Alpine treelines
- definitions vary widely - timberline can be considered to be the upper limit of closed forest, treeline can be considered to be a line connecting the highest patches of forest, and subalpine the transition zone
- most generally in the temperate zone, treeline changes about 75 m. per degree latitude
- less moisture, more sunshine and less exposure to frontal weather causes temperature in the central ranges to be higher and treeline to rise - the 'Massenerhebungseffect"
- increasing precipitation and cloudiness tends to suppress treelines
- treelines do not follow climatic change very rapidly - their current position must be assumed to reflect an environmental integral over several hundreds of years
- the author hypothesizes that there is a minimum mean temperature that permits sufficient production of new cells and the development and differentiation of functional tissue
- studies showing the diminishing growth rate with altitude support this concept
Chapter 8 - Climatic stress
- alpines have adapted to tolerate up to 10 degrees of freezing
- some plants have tissues adapted to allow supercooling so that nucleation does not occur until well below freezing
- many alpines are tolerant of herbivore browsing, showing that in most situations alpines are not carbon limited
- leaf temperatures of 50 degrees C are common in full sun
- flavanoids protect alpine plants from the higher level of UV radiation found at altitude
- much cell division and differentiation occurs below ground, protected from cold and heat
Chapter 9 - Water relations
- due to extensive root systems, soil moisture tends not to be limiting in the Alpine
- stomata density tends to increase with altitude, offsetting the decreased pressure; more stomata are found on the top of the leaf especially in cushion plants such as Saxifraga oppositifolia
- alpines have been found to open stomata for evaporative demand, not water supply
- as altitude increases, there is shift from C4 species to C3 species
- alpines have the ability to rehydrate after extensive periods of desiccation
- desiccation stress in higher plants is a very rare phenomenon in the alpine life zone
Chapter 10 - Mineral nutrition
- there are no indications of mineral shortages in alpine plants, showing that nutrition is not a limitation on plant size or altitude
- plants depend heavily on nitrogen recycling, compared with fixation
- alpine herbivores make a significant contribution to nitrogen recycling, as do grasshoppers
- mycorrhizal activity declines with altitude, becoming rare in isolated plants on high peaks
- plant distribution is heavily influenced by the supply of calcium
Chapter 11 - Uptake and loss of carbon
- the photosynthetic capacity of alpines varies by type, being 8 times higher in shrubs than lichens and twice a high in forbs than shrubs
- alpine plants have an extra cell layer in the leaves, making them thicker, and to some degree offsetting the decreased partial pressure of CO2 at higher altitudes
- high elevation plants fix carbon more efficiently (/m2) than lowland plants, apparently due to the differences in leaf anatomy
- photosynthesis in alpines is not restricted by temperature as high sunlight intensity raises leaf temperatures to optimal levels
- alpines in sunny locations are able to fix about five times the carbon actually contained in their leaves over the season
- plants studied in sunny locations are able to recoup their investment in leaves in 37 days of the 120 day season; shady plants required 68 days
- plants utilizing C4 photosynthesis are fewer as the elevation increases and are not generally found above treeline
- CAM plants tend to grow in drier locations; notable examples are cacti in South America
- alpines in their habitat respire at a rate similar to lowland plants
- some alpines do not adjust their respiration to higher temperatures and so cannot acclimatize to lower elevations
Chapter 12 - Carbon investments
- alpines accumulate large reserves of non-structural carbohydrates and of lipids, much of which is lost to the leaf litter; this apparent waste of carbon reserves is not understood
- alpines tend to develop less leaf area but greater root length than lowland species - root length may be due to reduced mycorrhizal activity at altitude
Chapter 13 - Growth dynamics and phenology
- the onset of spring growth depends on a specific photoperiod, after which snow cover and weather dictate
- senescence and the formation of winter buds are tightly tied to photoperiod
Chapter 14 - Cell division and tissue formation
- alpine plants produce cells similar in size to lowland plants, so their smaller size is due to fewer cells
- little is known of the mechanics of cell division in alpine plants
- "Perhaps we should then realize that small size is also part of a plan, a plan for survival and fitness, rather than the result of immediate limitations of a hostile environment"
Chapter 15 - Plant biomass production
- alpine plants are as efficient in energy conversion as other temperate region plants
- alpine herbivores include grasshoppers (Alps only?), guanacos in the Andes, and rodents - studies show that alpine plants have adapted to the continual grazing
Chapter 16 - Plant reproduction
- early flowering plants are opportunistic, flowering when the snow leaves, while late flowering plants do so only when the leaf crop has been produced, triggered by photoperiod
- in early plants, future years flowers are initiated as plants enter the winter; Polygonum viviparum requires three seasons of preparation to flower in the fourth year
- early plants have a lower reproductive success rate, but any seeds have a high chance of maturing, while late plants have a higher output of seeds which have a lower chance of maturing
- early plants suffer from a lack of pollinator activity, while late plants face competition for pollinators
- early plants have high outbreeding rates, while later plants are more given to selfing, apomixis and vivipary
- alpines tend to have flowers similar in size to lowland plants, but the flowering stalks are much reduced
- germination of alpines tend to require a winter quiescence, followed by warm temperatures
- clonal propagation is common in various alpine species as an alternate form of reproduction; forms such as stolons are expensive in terms of biomass
- in many species that engage in both sexual and clonal reproduction, the production of seed can be expensive and of low success but guarantees high genetic diversity and rapid colonization of new ground
- the age of clonal and tap rooted alpine forbs is frequently in the range of 30 - 50 years; slower growing species including cushion plants may be over 100 years of age; creeping shrubs "may be functionally immortal"
Chapter 17 - Global change at high elevation
- direct human influences on the alpine environment are a greater and more immediate concern than atmospheric changes
Chapter 1 - Plant ecology at high elevations
- conditions in the alpine provide natural experiments in comparative ecological research
Chapter 2 - The alpine life zone
- higher alpine plants grow up to 6000 m. in the Andes and Himalayas
- total alpine flora of the world is likely 8,000 to 10,000 higher plants
- alpine plants are typically prostrate woody shrubs, graminoids, cushion plants and herbaceous perennials often forming rosettes
- annuals commonly do not represent more than 2 percent of the alpine flora
Chapter 3 - Alpine climate
- while the lapse rate is usually taken to be 1 degree C per 100m, 0.6 degrees seems to be more typical for temperate summers and the tropics
- alpine plants can be subject to high radiation, both direct and diffuse from neighbouring clouds
Chapter 4 - The climate plants experience
- while conditions can be harsh in the alpine, it is different in the plant leaf canopy close to the ground where conditions may be 27 degrees C and 98 percent RH
- snow disappearance can vary with wind and exposure, varying by as much as 4 months - some plants can miss out one or two complete summers due to snow pack
- rosette plants can concentrate solar radiation, producing leaf temperatures over 50 degrees C
- Silene acaulis can produce leaf temperatures over 10 C degrees above air temperature; cushion plants become more abundant in windy environments
- similarly, tussock grasses use dead but sanding leaves to minimize wind effects
- root temperatures are low, often being only a few degrees above freezing
Chapter 5 - Life under snow
- alpines are able to photosynthesize at temperatures near freezing; some plants can reach 20 - 30 percent of their maximal rates
- as snow if not entirely reflective, plants are able to achieve net photosynthetic carbon gains under shallow snow
- even at 2 m. depth, photo signals are strong enough to induce seed germination, and sprouting
- CO2 concentrations under snow can exceed 1000 ppm, but this is in line with soil concentrations and do not appear to provide a significant benefit
- while oxygen concentration in snowpacks may be reduced, it does not appear to interfere with plant respiration
- the red colour of early leaves and stems is due to a lack of green chloroplasts allowing them to establish a radiation protective screen before the photosynthetic machinery is installed
- faster developing species prefer the center of snowbeds, while those taking more time prefer the edges
- it seems unlikely that sub-snow photosynthesis contributes significantly to the annual carbon balance of higher plants exposed to long-lasting snowpacks
Chapter 6 - Alpine soils
- dust accumulation has been found to have formed 10 cm of fine soil over the post-glacial period
- freezing is important in the heaving of bulk soils, resulting in mixing
- freeze-thaw causes downslope movement or solifluction, causing mechanical stress on the root systems
- accumulation of dead plant material results in biologically driven soil formation; essentially private recycling by cushion plants
- decomposition of leaf remains may take from 2 years for forbs to 5 years for sedges
- in the Rocky Mountains, it takes about 10,000 years for alpine soils to reach 30 cm
- the diversity of soil fungi decreases with altitude
Chapter 7 - Alpine treelines
- definitions vary widely - timberline can be considered to be the upper limit of closed forest, treeline can be considered to be a line connecting the highest patches of forest, and subalpine the transition zone
- most generally in the temperate zone, treeline changes about 75 m. per degree latitude
- less moisture, more sunshine and less exposure to frontal weather causes temperature in the central ranges to be higher and treeline to rise - the 'Massenerhebungseffect"
- increasing precipitation and cloudiness tends to suppress treelines
- treelines do not follow climatic change very rapidly - their current position must be assumed to reflect an environmental integral over several hundreds of years
- the author hypothesizes that there is a minimum mean temperature that permits sufficient production of new cells and the development and differentiation of functional tissue
- studies showing the diminishing growth rate with altitude support this concept
Chapter 8 - Climatic stress
- alpines have adapted to tolerate up to 10 degrees of freezing
- some plants have tissues adapted to allow supercooling so that nucleation does not occur until well below freezing
- many alpines are tolerant of herbivore browsing, showing that in most situations alpines are not carbon limited
- leaf temperatures of 50 degrees C are common in full sun
- flavanoids protect alpine plants from the higher level of UV radiation found at altitude
- much cell division and differentiation occurs below ground, protected from cold and heat
Chapter 9 - Water relations
- due to extensive root systems, soil moisture tends not to be limiting in the Alpine
- stomata density tends to increase with altitude, offsetting the decreased pressure; more stomata are found on the top of the leaf especially in cushion plants such as Saxifraga oppositifolia
- alpines have been found to open stomata for evaporative demand, not water supply
- as altitude increases, there is shift from C4 species to C3 species
- alpines have the ability to rehydrate after extensive periods of desiccation
- desiccation stress in higher plants is a very rare phenomenon in the alpine life zone
Chapter 10 - Mineral nutrition
- there are no indications of mineral shortages in alpine plants, showing that nutrition is not a limitation on plant size or altitude
- plants depend heavily on nitrogen recycling, compared with fixation
- alpine herbivores make a significant contribution to nitrogen recycling, as do grasshoppers
- mycorrhizal activity declines with altitude, becoming rare in isolated plants on high peaks
- plant distribution is heavily influenced by the supply of calcium
Chapter 11 - Uptake and loss of carbon
- the photosynthetic capacity of alpines varies by type, being 8 times higher in shrubs than lichens and twice a high in forbs than shrubs
- alpine plants have an extra cell layer in the leaves, making them thicker, and to some degree offsetting the decreased partial pressure of CO2 at higher altitudes
- high elevation plants fix carbon more efficiently (/m2) than lowland plants, apparently due to the differences in leaf anatomy
- photosynthesis in alpines is not restricted by temperature as high sunlight intensity raises leaf temperatures to optimal levels
- alpines in sunny locations are able to fix about five times the carbon actually contained in their leaves over the season
- plants studied in sunny locations are able to recoup their investment in leaves in 37 days of the 120 day season; shady plants required 68 days
- plants utilizing C4 photosynthesis are fewer as the elevation increases and are not generally found above treeline
- CAM plants tend to grow in drier locations; notable examples are cacti in South America
- alpines in their habitat respire at a rate similar to lowland plants
- some alpines do not adjust their respiration to higher temperatures and so cannot acclimatize to lower elevations
Chapter 12 - Carbon investments
- alpines accumulate large reserves of non-structural carbohydrates and of lipids, much of which is lost to the leaf litter; this apparent waste of carbon reserves is not understood
- alpines tend to develop less leaf area but greater root length than lowland species - root length may be due to reduced mycorrhizal activity at altitude
Chapter 13 - Growth dynamics and phenology
- the onset of spring growth depends on a specific photoperiod, after which snow cover and weather dictate
- senescence and the formation of winter buds are tightly tied to photoperiod
Chapter 14 - Cell division and tissue formation
- alpine plants produce cells similar in size to lowland plants, so their smaller size is due to fewer cells
- little is known of the mechanics of cell division in alpine plants
- "Perhaps we should then realize that small size is also part of a plan, a plan for survival and fitness, rather than the result of immediate limitations of a hostile environment"
Chapter 15 - Plant biomass production
- alpine plants are as efficient in energy conversion as other temperate region plants
- alpine herbivores include grasshoppers (Alps only?), guanacos in the Andes, and rodents - studies show that alpine plants have adapted to the continual grazing
Chapter 16 - Plant reproduction
- early flowering plants are opportunistic, flowering when the snow leaves, while late flowering plants do so only when the leaf crop has been produced, triggered by photoperiod
- in early plants, future years flowers are initiated as plants enter the winter; Polygonum viviparum requires three seasons of preparation to flower in the fourth year
- early plants have a lower reproductive success rate, but any seeds have a high chance of maturing, while late plants have a higher output of seeds which have a lower chance of maturing
- early plants suffer from a lack of pollinator activity, while late plants face competition for pollinators
- early plants have high outbreeding rates, while later plants are more given to selfing, apomixis and vivipary
- alpines tend to have flowers similar in size to lowland plants, but the flowering stalks are much reduced
- germination of alpines tend to require a winter quiescence, followed by warm temperatures
- clonal propagation is common in various alpine species as an alternate form of reproduction; forms such as stolons are expensive in terms of biomass
- in many species that engage in both sexual and clonal reproduction, the production of seed can be expensive and of low success but guarantees high genetic diversity and rapid colonization of new ground
- the age of clonal and tap rooted alpine forbs is frequently in the range of 30 - 50 years; slower growing species including cushion plants may be over 100 years of age; creeping shrubs "may be functionally immortal"
Chapter 17 - Global change at high elevation
- direct human influences on the alpine environment are a greater and more immediate concern than atmospheric changes
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