@article{penuelasBVOCsPlantDefense2015,
  title = {{{BVOCs}}: Plant Defense against Climate Warming?},
  author = {Pe{\~n}uelas, Josep and Llusi{\`a}, Joan},
  year = {2015},
  month = jun,
  volume = {8},
  pages = {105--109},
  doi = {10.1016/s1360-1385(03)00008-6},
  abstract = {Plants emit a substantial amount of biogenic volatile organic compounds (BVOCs) into the atmosphere. These BVOCs represent a large carbon loss and can be up to {$\sim$}10\,\% of that fixed by photosynthesis under stressful conditions and up to 100 g C m-2 per year in some tropical ecosystems. Among a variety of proven and unproven BVOC functions in plants and roles in atmospheric processes, recent data intriguingly link emission of these compounds to climate. Ongoing research demonstrates that BVOCs could protect plants against high temperatures. BVOC emissions are probably increasing with warming and with other factors associated to global change, including changes in land cover. These increases in BVOC emissions could contribute in a significant way (via negative and positive feedback) to the complex processes associated with global warming.

[\textbackslash n] Plants re-emit a substantial fraction of their assimilated carbon into the atmosphere as biogenic volatile organic compounds (BVOCs) that affect the chemical and physical properties of the atmosphere 1, 2, 3 and 4. Why do plants emit them? What are their effects on atmospheric chemistry and physics? These are two crucial questions that have been challenging the scientific community for several years. Among a variety of likely BVOCs functions in plants and effects on the atmosphere, recent data intriguingly link these compounds with climate. BVOCs could protect plants against high temperatures. But, in their turn, BVOC emissions increase with warming, and might produce both negative and positive feedback on climate warming through aerosol formation and direct and indirect greenhouse effects.

[Excerpt] 

[::Nature and function of BVOCs] BVOCs are produced in many different plant tissues and physiological processes. They are diverse, including isoprene, terpenes, alkanes, alkenes, alcohols, esters, carbonyls and acids (Table 1). Indeed, the enormous variety of BVOCs constitutes one of nature's biodiversity treasures. Advances in molecular and genetic techniques and development of new instrumentation for the collection and analysis of BVOCs have increased our knowledge of their nature and function in recent years. In some plants, BVOCs accumulate in specialized organs in leaves and stems and can be released as deterrents against pathogens and herbivores, or to aid wound sealing after damage [5]. In other plants, BVOCs are not stored and are emitted after production. The purpose of BVOCs appears to be to attract pollinators and herbivore predators, and to communicate with other plants and organisms 6 and 7. But there is another function of BVOCs that has attracted interest given current climate warming. Recently, some evidence has emerged that the production and the emission of BVOCs, such as isoprene and monoterpenes, which constitute a major fraction of BVOCs, might confer protection against high temperatures.

[::Thermotolerance and photorespiration]

Thomas Sharkey and Eric Singsaas [8] were the first people to propose that isoprene has a thermotolerance function. Later, thermotolerance was also observed in monoterpene emissions from Quercus ilex, a Mediterranean oak species [9]. Further studies have not always been able to reproduce these results and the mechanism for this protection against high-temperature damage has yet to be shown [10]. However, recent research shows a possible link with photorespiration [11], another relatively poorly understood plant process. Earlier work had already suggested that isoprene biosynthesis was linked to photorespiration [12]. However, subsequent results with Populus tremuloides [13], or Arundo donax [14], among others, did not find such a relationship. The most recent studies have not just focused on photorespiration as an alternative source of carbon for isoprenoid biosynthesis when photosynthesis is limited. They have shown that the formation of monoterpenes might depend on photorespiratory activity, and that under non-photorespiratory conditions monoterpenes seem to replace photorespiration in providing protection against high temperatures [11]. Therefore, in agreement with previous suggestions of isoprene as a scavenger of hydroxyl radicals and as a protector of membranes and macromolecules from oxidative damage [15], it is likely that monoterpenes protect plant tissues as scavengers of reactive oxygen species produced under high temperatures, especially when photorespiration is not active in inhibiting their formation. This protection would not be surprising because isoprene and monoterpenes would have, thus, a parallel function to that of larger isoprenoids such as xanthophylls [16]. [...]

[::Conclusions and open questions]

In conclusion, among the great variety of likely BVOC functions in plants and effects on the atmosphere, recent data intriguingly link these compounds with climate. BVOCs could protect plants against high temperatures in a process linked to photorespiration. But, in turn, BVOCs emissions increase with warming and with most of the other components of the current global environmental change. And this increase, apart from influencing the oxidizing potential of the troposphere, might produce both negative and positive feedback on warming depending on the spatial scales, and on the relative effects of aerosol formation and direct and indirect greenhouse properties.

[\textbackslash n] Many questions about these BVOC relationships with both plant and atmosphere temperatures remain to be solved. Regarding plant thermoprotection: do plants always protect themselves from over-heating by producing and emitting BVOCs? Are there species characteristics, physiological states or environmental conditions that determine this photoprotection? How does photorespiration affect monoterpene synthesis? Which are the mechanisms and the interactions involved? Are BVOCs just scavengers of radical oxygen species or can they also act as membrane stabilizers [40]? To what extent might BVOC emissions directly cool the plant through physiological or evaporative effects?

[\textbackslash n] There are also many unanswered questions regarding the relationship between BVOCs and global change, but they can be summarized by asking: how much will BVOC emissions increase in response to the global change drivers? The scientific community needs to focus not only on the climate-warming effects on BVOC emissions, but also on the effects of the other global change drivers, especially on the increasing land-cover changes. The species- and ecosystem-specificity of BVOC emissions and the possible role of plant species and ecosystems as sinks of BVOCs should also be taken into account.

[\textbackslash n] And finally, regarding the climate relationships, whether the increased BVOC emissions will cool or warm the environment needs to be determined. The overall effect of increasing BVOC emissions will depend on the relative weight and scale of the negative (increased albedo) and positive (increased greenhouse action) feedback. As is the case with so many environmental issues, interactive interdisciplinary research among biologists, physicists and chemists at foliar, ecosystem, regional and global scales is needed to solve these puzzles.},
  journal = {Trends in Plant Science},
  keywords = {*imported-from-citeulike-INRMM,~INRMM-MiD:c-13646943,~to-add-doi-URL,biogenic-volatile-organic-compounds,carbon-cycle,climate,complexity,ecosystem-services,feedback,forest-resources,global-warming,homeostasis,isoprene,monoterpenes,off-site-effects,quercus-ilex},
  lccn = {INRMM-MiD:c-13646943},
  number = {3}
}

{"_id":"EgxMKoZ8acLRHQcDZ","bibbaseid":"peuelas-llusi-bvocsplantdefenseagainstclimatewarming-2015","downloads":0,"creationDate":"2016-06-22T10:19:42.305Z","title":"BVOCs: Plant Defense against Climate Warming?","author_short":["Peñuelas, J.","Llusià, J."],"year":2015,"bibtype":"article","biburl":"https://sharefast.me/php/download.php?id=zOUKvA&token=29","bibdata":{"bibtype":"article","type":"article","title":"BVOCs: Plant Defense against Climate Warming?","author":[{"propositions":[],"lastnames":["Peñuelas"],"firstnames":["Josep"],"suffixes":[]},{"propositions":[],"lastnames":["Llusià"],"firstnames":["Joan"],"suffixes":[]}],"year":"2015","month":"June","volume":"8","pages":"105–109","doi":"10.1016/s1360-1385(03)00008-6","abstract":"Plants emit a substantial amount of biogenic volatile organic compounds (BVOCs) into the atmosphere. These BVOCs represent a large carbon loss and can be up to $\\sim$10\\,% of that fixed by photosynthesis under stressful conditions and up to 100 g C m-2 per year in some tropical ecosystems. Among a variety of proven and unproven BVOC functions in plants and roles in atmospheric processes, recent data intriguingly link emission of these compounds to climate. Ongoing research demonstrates that BVOCs could protect plants against high temperatures. BVOC emissions are probably increasing with warming and with other factors associated to global change, including changes in land cover. These increases in BVOC emissions could contribute in a significant way (via negative and positive feedback) to the complex processes associated with global warming. [\\n] Plants re-emit a substantial fraction of their assimilated carbon into the atmosphere as biogenic volatile organic compounds (BVOCs) that affect the chemical and physical properties of the atmosphere 1, 2, 3 and 4. Why do plants emit them? What are their effects on atmospheric chemistry and physics? These are two crucial questions that have been challenging the scientific community for several years. Among a variety of likely BVOCs functions in plants and effects on the atmosphere, recent data intriguingly link these compounds with climate. BVOCs could protect plants against high temperatures. But, in their turn, BVOC emissions increase with warming, and might produce both negative and positive feedback on climate warming through aerosol formation and direct and indirect greenhouse effects. [Excerpt] [::Nature and function of BVOCs] BVOCs are produced in many different plant tissues and physiological processes. They are diverse, including isoprene, terpenes, alkanes, alkenes, alcohols, esters, carbonyls and acids (Table 1). Indeed, the enormous variety of BVOCs constitutes one of nature's biodiversity treasures. Advances in molecular and genetic techniques and development of new instrumentation for the collection and analysis of BVOCs have increased our knowledge of their nature and function in recent years. In some plants, BVOCs accumulate in specialized organs in leaves and stems and can be released as deterrents against pathogens and herbivores, or to aid wound sealing after damage [5]. In other plants, BVOCs are not stored and are emitted after production. The purpose of BVOCs appears to be to attract pollinators and herbivore predators, and to communicate with other plants and organisms 6 and 7. But there is another function of BVOCs that has attracted interest given current climate warming. Recently, some evidence has emerged that the production and the emission of BVOCs, such as isoprene and monoterpenes, which constitute a major fraction of BVOCs, might confer protection against high temperatures. [::Thermotolerance and photorespiration] Thomas Sharkey and Eric Singsaas [8] were the first people to propose that isoprene has a thermotolerance function. Later, thermotolerance was also observed in monoterpene emissions from Quercus ilex, a Mediterranean oak species [9]. Further studies have not always been able to reproduce these results and the mechanism for this protection against high-temperature damage has yet to be shown [10]. However, recent research shows a possible link with photorespiration [11], another relatively poorly understood plant process. Earlier work had already suggested that isoprene biosynthesis was linked to photorespiration [12]. However, subsequent results with Populus tremuloides [13], or Arundo donax [14], among others, did not find such a relationship. The most recent studies have not just focused on photorespiration as an alternative source of carbon for isoprenoid biosynthesis when photosynthesis is limited. They have shown that the formation of monoterpenes might depend on photorespiratory activity, and that under non-photorespiratory conditions monoterpenes seem to replace photorespiration in providing protection against high temperatures [11]. Therefore, in agreement with previous suggestions of isoprene as a scavenger of hydroxyl radicals and as a protector of membranes and macromolecules from oxidative damage [15], it is likely that monoterpenes protect plant tissues as scavengers of reactive oxygen species produced under high temperatures, especially when photorespiration is not active in inhibiting their formation. This protection would not be surprising because isoprene and monoterpenes would have, thus, a parallel function to that of larger isoprenoids such as xanthophylls [16]. [...] [::Conclusions and open questions] In conclusion, among the great variety of likely BVOC functions in plants and effects on the atmosphere, recent data intriguingly link these compounds with climate. BVOCs could protect plants against high temperatures in a process linked to photorespiration. But, in turn, BVOCs emissions increase with warming and with most of the other components of the current global environmental change. And this increase, apart from influencing the oxidizing potential of the troposphere, might produce both negative and positive feedback on warming depending on the spatial scales, and on the relative effects of aerosol formation and direct and indirect greenhouse properties. [\\n] Many questions about these BVOC relationships with both plant and atmosphere temperatures remain to be solved. Regarding plant thermoprotection: do plants always protect themselves from over-heating by producing and emitting BVOCs? Are there species characteristics, physiological states or environmental conditions that determine this photoprotection? How does photorespiration affect monoterpene synthesis? Which are the mechanisms and the interactions involved? Are BVOCs just scavengers of radical oxygen species or can they also act as membrane stabilizers [40]? To what extent might BVOC emissions directly cool the plant through physiological or evaporative effects? [\\n] There are also many unanswered questions regarding the relationship between BVOCs and global change, but they can be summarized by asking: how much will BVOC emissions increase in response to the global change drivers? The scientific community needs to focus not only on the climate-warming effects on BVOC emissions, but also on the effects of the other global change drivers, especially on the increasing land-cover changes. The species- and ecosystem-specificity of BVOC emissions and the possible role of plant species and ecosystems as sinks of BVOCs should also be taken into account. [\\n] And finally, regarding the climate relationships, whether the increased BVOC emissions will cool or warm the environment needs to be determined. The overall effect of increasing BVOC emissions will depend on the relative weight and scale of the negative (increased albedo) and positive (increased greenhouse action) feedback. As is the case with so many environmental issues, interactive interdisciplinary research among biologists, physicists and chemists at foliar, ecosystem, regional and global scales is needed to solve these puzzles.","journal":"Trends in Plant Science","keywords":"*imported-from-citeulike-INRMM,~INRMM-MiD:c-13646943,~to-add-doi-URL,biogenic-volatile-organic-compounds,carbon-cycle,climate,complexity,ecosystem-services,feedback,forest-resources,global-warming,homeostasis,isoprene,monoterpenes,off-site-effects,quercus-ilex","lccn":"INRMM-MiD:c-13646943","number":"3","bibtex":"@article{penuelasBVOCsPlantDefense2015,\n title = {{{BVOCs}}: Plant Defense against Climate Warming?},\n author = {Pe{\\~n}uelas, Josep and Llusi{\\`a}, Joan},\n year = {2015},\n month = jun,\n volume = {8},\n pages = {105--109},\n doi = {10.1016/s1360-1385(03)00008-6},\n abstract = {Plants emit a substantial amount of biogenic volatile organic compounds (BVOCs) into the atmosphere. These BVOCs represent a large carbon loss and can be up to {$\\sim$}10\\,\\% of that fixed by photosynthesis under stressful conditions and up to 100 g C m-2 per year in some tropical ecosystems. Among a variety of proven and unproven BVOC functions in plants and roles in atmospheric processes, recent data intriguingly link emission of these compounds to climate. Ongoing research demonstrates that BVOCs could protect plants against high temperatures. BVOC emissions are probably increasing with warming and with other factors associated to global change, including changes in land cover. These increases in BVOC emissions could contribute in a significant way (via negative and positive feedback) to the complex processes associated with global warming.\n\n[\\textbackslash n] Plants re-emit a substantial fraction of their assimilated carbon into the atmosphere as biogenic volatile organic compounds (BVOCs) that affect the chemical and physical properties of the atmosphere 1, 2, 3 and 4. Why do plants emit them? What are their effects on atmospheric chemistry and physics? These are two crucial questions that have been challenging the scientific community for several years. Among a variety of likely BVOCs functions in plants and effects on the atmosphere, recent data intriguingly link these compounds with climate. BVOCs could protect plants against high temperatures. But, in their turn, BVOC emissions increase with warming, and might produce both negative and positive feedback on climate warming through aerosol formation and direct and indirect greenhouse effects.\n\n[Excerpt] \n\n[::Nature and function of BVOCs] BVOCs are produced in many different plant tissues and physiological processes. They are diverse, including isoprene, terpenes, alkanes, alkenes, alcohols, esters, carbonyls and acids (Table 1). Indeed, the enormous variety of BVOCs constitutes one of nature's biodiversity treasures. Advances in molecular and genetic techniques and development of new instrumentation for the collection and analysis of BVOCs have increased our knowledge of their nature and function in recent years. In some plants, BVOCs accumulate in specialized organs in leaves and stems and can be released as deterrents against pathogens and herbivores, or to aid wound sealing after damage [5]. In other plants, BVOCs are not stored and are emitted after production. The purpose of BVOCs appears to be to attract pollinators and herbivore predators, and to communicate with other plants and organisms 6 and 7. But there is another function of BVOCs that has attracted interest given current climate warming. Recently, some evidence has emerged that the production and the emission of BVOCs, such as isoprene and monoterpenes, which constitute a major fraction of BVOCs, might confer protection against high temperatures.\n\n[::Thermotolerance and photorespiration]\n\nThomas Sharkey and Eric Singsaas [8] were the first people to propose that isoprene has a thermotolerance function. Later, thermotolerance was also observed in monoterpene emissions from Quercus ilex, a Mediterranean oak species [9]. Further studies have not always been able to reproduce these results and the mechanism for this protection against high-temperature damage has yet to be shown [10]. However, recent research shows a possible link with photorespiration [11], another relatively poorly understood plant process. Earlier work had already suggested that isoprene biosynthesis was linked to photorespiration [12]. However, subsequent results with Populus tremuloides [13], or Arundo donax [14], among others, did not find such a relationship. The most recent studies have not just focused on photorespiration as an alternative source of carbon for isoprenoid biosynthesis when photosynthesis is limited. They have shown that the formation of monoterpenes might depend on photorespiratory activity, and that under non-photorespiratory conditions monoterpenes seem to replace photorespiration in providing protection against high temperatures [11]. Therefore, in agreement with previous suggestions of isoprene as a scavenger of hydroxyl radicals and as a protector of membranes and macromolecules from oxidative damage [15], it is likely that monoterpenes protect plant tissues as scavengers of reactive oxygen species produced under high temperatures, especially when photorespiration is not active in inhibiting their formation. This protection would not be surprising because isoprene and monoterpenes would have, thus, a parallel function to that of larger isoprenoids such as xanthophylls [16]. [...]\n\n[::Conclusions and open questions]\n\nIn conclusion, among the great variety of likely BVOC functions in plants and effects on the atmosphere, recent data intriguingly link these compounds with climate. BVOCs could protect plants against high temperatures in a process linked to photorespiration. But, in turn, BVOCs emissions increase with warming and with most of the other components of the current global environmental change. And this increase, apart from influencing the oxidizing potential of the troposphere, might produce both negative and positive feedback on warming depending on the spatial scales, and on the relative effects of aerosol formation and direct and indirect greenhouse properties.\n\n[\\textbackslash n] Many questions about these BVOC relationships with both plant and atmosphere temperatures remain to be solved. Regarding plant thermoprotection: do plants always protect themselves from over-heating by producing and emitting BVOCs? Are there species characteristics, physiological states or environmental conditions that determine this photoprotection? How does photorespiration affect monoterpene synthesis? Which are the mechanisms and the interactions involved? Are BVOCs just scavengers of radical oxygen species or can they also act as membrane stabilizers [40]? To what extent might BVOC emissions directly cool the plant through physiological or evaporative effects?\n\n[\\textbackslash n] There are also many unanswered questions regarding the relationship between BVOCs and global change, but they can be summarized by asking: how much will BVOC emissions increase in response to the global change drivers? The scientific community needs to focus not only on the climate-warming effects on BVOC emissions, but also on the effects of the other global change drivers, especially on the increasing land-cover changes. The species- and ecosystem-specificity of BVOC emissions and the possible role of plant species and ecosystems as sinks of BVOCs should also be taken into account.\n\n[\\textbackslash n] And finally, regarding the climate relationships, whether the increased BVOC emissions will cool or warm the environment needs to be determined. The overall effect of increasing BVOC emissions will depend on the relative weight and scale of the negative (increased albedo) and positive (increased greenhouse action) feedback. As is the case with so many environmental issues, interactive interdisciplinary research among biologists, physicists and chemists at foliar, ecosystem, regional and global scales is needed to solve these puzzles.},\n journal = {Trends in Plant Science},\n keywords = {*imported-from-citeulike-INRMM,~INRMM-MiD:c-13646943,~to-add-doi-URL,biogenic-volatile-organic-compounds,carbon-cycle,climate,complexity,ecosystem-services,feedback,forest-resources,global-warming,homeostasis,isoprene,monoterpenes,off-site-effects,quercus-ilex},\n lccn = {INRMM-MiD:c-13646943},\n number = {3}\n}\n\n","author_short":["Peñuelas, J.","Llusià, J."],"key":"penuelasBVOCsPlantDefense2015","id":"penuelasBVOCsPlantDefense2015","bibbaseid":"peuelas-llusi-bvocsplantdefenseagainstclimatewarming-2015","role":"author","urls":{},"keyword":["*imported-from-citeulike-INRMM","~INRMM-MiD:c-13646943","~to-add-doi-URL","biogenic-volatile-organic-compounds","carbon-cycle","climate","complexity","ecosystem-services","feedback","forest-resources","global-warming","homeostasis","isoprene","monoterpenes","off-site-effects","quercus-ilex"],"downloads":0},"search_terms":["bvocs","plant","defense","against","climate","warming","peñuelas","llusià"],"keywords":["biogenic-volatile-organic-compounds","carbon-cycle","climate","complexity","ecosystem-services","feedback","forest-resources","global-warming","homeostasis","isoprene","monoterpenes","off-site-effects","quercus-ilex","*imported-from-citeulike-inrmm","~inrmm-mid:c-13646943","~to-add-doi-url"],"authorIDs":[],"dataSources":["5S2zj2hKW8TWTkuMq"]}