Effects of Enhanced CO2 on Tropical Forest Growth

Effects of Enhanced CO2 on Tropical Forest Growth James P. Smith Effects of enhanced atmospheric CO2 concentrations on tropical forest growth: experimental studies and interactions with nutrients, light, water and temperature Abstract (150 words) Introduction (300 words) Approximately 90% of earth’s 652Gt terrestrial biomass carbon is locked up in forests. Tropical and subtropical forests store 340Gt carbon; or ~52%; but only make up 13% of total forested area (table 1). Achard et al (2002) estimated 1Gt/yr carbon losses, through activities such as deforestation and clearance for agriculture (Geist et al, 2002). All terrestrial plants have become exposed to increasing atmospheric CO2 concentrations, as part of global change. This has changed from 180ppm 18ka (Petit et al, 1999) to 390ppm today, by degassing from oceans and fossil C burning (Crowley et al, 2001). Increased CO2 could stimulate photosynthesis, raising plant productivity. This can have a role in storing more carbon and mitigate the atmospheric rise in CO2 concentrations (Beedlow et al, 2004). Table 1: Areal extent, carbon storage and net primary productivity of earth’s major biomes (from Roy et al, 2001). Figure 1 demonstrates CO2 enters plants at the source (leaf); where it becomes photoassimilated to produce carbon sugars; which are transported around the plant to carbon sinks; for different processes such as structural growth, metabolism and export. Sugars can also be stored as reserves in the form of NSCs (non-structural carbohydrates). CO2 is lost through respiration, herbivory and litter production and decomposition (Korner, 2003a). Figure 1: CO2 pools and fluxes in plants, as well as source-sink interactions (modified from Korner, 2003a). The aim of the review is to evaluate research on the effects of enhanced CO2 on tropical forest growth. This will be achieved by looking at experimental studies, as well as the effects of enhanced CO2 on the limiting factors of nutrients, light, water supply and temperature. I will be reviewing literature from 1999-2013. Experimental studies There have been few experimental studies of the effects of enhanced CO2 on plant growth in tropical forests in relatively natural conditions (ambient climate, natural soil and inter and intra-species competition). Two studies using a canopy crane in a tropical dry forest in Panama was used to assess the effects of enhanced CO2 on canopy tree leaves. Over a 40 week period Lovelock et al (1999) measured responses of leaf and branchlets of a single tree species. Photosynthesis rates increased 30% with enhanced CO2. However, no increases in biomass occurred (reproductive organs and foliage). Branchlet TNC (total non-structural carbohydrates) increased 20%, inferring localized carbon saturation. Wurth et al (1998a) found stronger TNC increases (41-61%), upon exposing canopy leaves of four tree species to enhanced CO2, in situ. Wurth et al (1998b) planted seedlings of five local species (tree, shrubs and grass) in the understorey of a closed Panamanian forest. These were grown over a 15 mo nth period, in which 50% were in ambient CO2 and 50% in elevated. All species showed significant seedling growth under elevated CO2, but decreased as understorey light levels increased, and inter-species variation was apparent. Again TNC levels increased under enhanced CO2. One experiment has studied communities of tropical trees, which have been outplanted in natural soil and subjected to elevated CO2. Lovelock et al (1998) grew groups of ten tree species at ambient and elevated CO2 in open-top chambers at the forest margin in Panama. Over six months, there was no enhancement in biomass accumulation. There were also reductions in leaf area index, increased photosynthesis rates and increased nitrogen: carbon ratios. Response was species-specific, but late-successional species were less sensitive than pioneer and midsuccessional species. Table 2: Comparison of mean TNC concentrations (% dry weight) across four studies under ambient and elevated CO2 concentrations. From table 2, it is clear that all four studies mentioned showed increased mean TNC concentrations when exposed to elevated CO2. Despite the increases, this does not necessarily mean TNCs from carbon sources are being transported to carbon sinks, into plant biomass for growth. They include carbohydrates, sugar alcohols, organic acids and lipids, and represents carbon reserves or stores, for future use on demand (Korner, 2003a). So, photosynthesis rates may increase under elevated CO2, producing more TNCs, but may not be used in plant growth, unless needed. Figure 2: Variation in mean concentration of TNC with height in two wet and dry seasons (from Wurth et al, 1998a). Wurth et al (1998a) also compared TNC concentrations, exposed to elevated CO2, with height from canopy height to roots, between wet and dry seasons (figure 2). They found TNC to increase in all plant compartments during the dry season. The TNC again not incorporated into structural growth, because growth was directly limited by dry conditions, and not photosynthesis. More TNC was being stored in reserves. In the wet season, TNC pools reduced, coinciding with resumed tree growth and new leaf production. They inferred TNC concentrations were controlled by moisture availability, in agreement with another study in the area (Newell et al, 2002). On the other hand, Korner and Wurth (1996) found TNC to increase significantly in both dry and wet seasons. This infers plants have a store of carbon, and can mobilize it when needed for growth. To further the understanding of increasing CO2 on tropical forest growth, more and longer-term experiments are needed. Arnone (1996) and Korner (1998) criticize these experiments, as they cannot be scaled up to actual forest size; use only small plants; have a higher than normal nutrient supply; absence of competition; and key processes; such as herbivory and effects of pathogens. CO2-nutrient interactions Nitrogen is commonly seen as the main limiting nutrient of tree CO2 responses (Finzi et al, 2006). However, although this is theoretically an unlimited resource (atmospheric), provided N fixation balances N losses through processes such as N20 losses or leaching (Korner, 2009). Litter mineralization is the predominate source of N in forests. All other nutrients are in limited supply in a given area, with older, more weathered (humid tropics) soils making these nutrients much more limiting to plant growth (Bergametti et al, 1998). Enhanced CO2 can accelerate the rate of symbiotic N fixation, as demonstrated by Tissue et al (1997). Seeds of fast-growing woody legumes from a seasonal tropical forest in Costa Rica were inoculated with N2 fixing Rhizobium bacteria and grown in greenhouses for ~70 days, exposed to ambient (35Pa) and elevated (70Pa) CO2 levels. Seedlings were watered adequately with N-free water solution. Under elevated CO2, photosynthesis rates increased by 49%, compared to those exposed to ambient CO2. As a result growth in elevated CO2 increased 36%. Figure 3 illustrates this, with total plant biomass growing 84% under elevated CO2. Greater rates of photosynthesis mean greater quantities of carbon are transported to the nodules. More carbon supplied to nodules means specific nitrogenase activity (SNA); that is N-fixing enzyme activity; is increased; more energy is available to power the fixation process. Thus a greater proportion of nitrogen is fixed by the legumes and incorporated into the plant for biomass accumulation and growth. Figure 4 shows this clearly, with increases in N content across all parts of the plant. Figures 3 4: Dry weight biomass (gDW) of whole plant, as well as different areas of the plant (left). N content (mg) of whole plant, and different sections of plant (right). (From Tissue et al, 1997). Although there is a high abundance of nitrogen, and fixing increases under CO2 levels, Pons et al (2007) inferred N-fixation is also strongly limited by phosphorus availability, and is absorbed by trees much more efficiently than N (Medina and Cuevas, 1994; Herbert and Fownes, 1995). Pons et al (2007) measured N and P concentration changes in leaves of leguminous plants, in different soil types, in a tropical forest in Guyana. From table 3, general increases in N and P led to positive accumulations of N in leaves. They inferred increases in phosphorus were the main cause for increasing N-fixation, with increasing N concentrations having negligible effect. Contrary to Tissue et al (1997)’s findings, Houlton et al (2008) found N fixation to be less prominent in tropical forests. Pons et al (2007) approximated 6% of total N uptake by trees in Guyana was by N-fixation, and only ~50% legumes used the symbiotic pathway. Nardoto et al (2008) found near negligible N-fixation levels in legumes in Amazonia. Thus, nitrogen is unlikely to majorly constrain C-fixation in tropical forests, but phosphorus is more likely to (Martinelli et al, 1999). Table 3: Phosphorus and nitrogen concentrations in five different soil types, and their affect on N-fixation rates by N contents in leaves (Modified from Pons et al, 2007). Studies in tropical forests in Panama provided clear evidence that trees grown in close proximity to their natural habitat, under elevated CO2, within original soils and under local climatic conditions, exhibited accelerated growth rates when soils were enriched with mineral nutrients (Winter and Lovelock, 1999; Winter et al, 2001; table 4). In the absence of fertilizer there was no significant change in growth rate under elevated CO2 (Lovelock et al, 1998; Winter et al, 2000). No major changes in growth rates were found again were found by Korner and Arnone (1992) and Arnone and Korner (1995). Table 4: The effect of fertilizer/absence of fertilizer application on biomass accumulation for tropical plants under elevated CO2. Clearly the effects of elevated CO2 on have caused mixed responses from different studies. In some studies, greater photosynthesis rates led to increased carbon supply to allow accelerated N-fixation for biomass growth. Other studies highlighted the greater importance of phosphorus in regulating N-fixation and biomass accumulation. Plants grown in the absence of nutrients consistently showed minimal to no change in growth rates, opposed to increasing biomass with those that were enriched with mineral nutrients. CO2-light interactions It is known that shaded plant growth rates are limited by light and CO2. Illuminating plants will lead to accelerated growth, by forest canopy thinning or removal. As enhanced CO2 increases light use efficiency and decreases the light compensation point within the leaf, stimulation by enhanced CO2 in shaded areas can be seen to be similar to canopy thinning or illumination (Long and Drake, 1991). The effect of elevated CO2 on tropical plants grown in deep shade can be significant and can possibly exceed effects grown under horticultural conditions under full light (Korner, 2009). Wurth (1998a) exposed seedlings on the forest floor to ~700ppm CO2 under extremely low light levels (~11ÃŽ ¼mol photons m-2s-1). Tree seedlings grew 25-44% and shrub seedlings grew 59-76%. Lovelock et al (1996) observed similar results of mycorrhizal growth of tree seedlings, although P supply may have had an influence. Thus elevated CO2 promotes expansion into shaded areas. As expressed, as most tree seedlings wait to exploit an opening in the canopy, lianas employ a different strategy. Lianas are situated in deep shade and aim to occupy maximal space, but with minimal structural investment (Korner, 2009). Elevated CO2 increases the probability of lianas reaching the upper canopy. Granados and Korner (2002) studied biomass and growth rates for three liana species; simulated in a tropical understorey environment with seed and soil from Yucatan; under high and low light levels; and under ambient and elevated CO2 levels. From figures 5-7 it is apparent that liana biomass increases at higher light levels for all three species. However, liana growth rate is much larger at lower light levels (up to +249%), opposed to higher light levels (up to +52%). These higher growth rates are at moderately elevated CO2 levels of ~420ppm. At ~700ppm, growth rates reduced or even reversed. Thus, individuals within the understorey with low light levels (under moderately elevated CO2 levels) have the potential to grow upwards towards the canopy at a faster rate than those in higher light levels. Figure 8: Comparison of biomass change and growth rates; under ambient and elevated CO2 concentrations; between temperate and tropical liana species (from Korner, 2009) This consistent trend in increased growth rates under low light levels has also been confirmed for temperate liana species (figure 8). Hattenschweiler and Korner (2003) found growth rates between 64-80% under low light opposed to 23-40% under high light. These results could support reasoning for the enhanced vigour and reproduction of lianas observed in recent decades in Panama (Wright et al, 2004) and Amazonia (Phillips et al, 2002). Elevated CO2 may cause lianas to behave more aggressively, thereby inducing faster forest turnover, and reducing tree carbon storage in the long-run (Korner, 2004). Other factors have also been attributed to explain current liana growth, such as reduced rainfall (Swaine and Grace, 2007). Epiphytes are another important organism that influence tropical forest tree dynamics, and grow in tree crowns. Epiphytes derive from succulents, and may utilize CAM (Crassulacean acid metabolism) photosynthetic pathways, although some can use C3 pathways also (Korner, 2009). Contrary to lianas, evidence suggests epiphytes don’t benefit from elevated CO2 (Monterio et al, 2009). They tested the effect of doubling CO2 concentration; as well as increasing light and nutrient levels; on growth of six epiphyte species from the Neotropics. Figure 9: Relative growth rate (mgg-1d-1) of six epiphyte species under increasing CO2, light and nutrient levels for six different species. C3 pathways (V=Vriesea; C=Catopsis; O=Oncidium). CAM pathways (T=Tillandsia; B=Bulbophyllum; A=Aechmea). From Monteiro et al (2009). From figure 9; across the six species; elevated CO2 increased relative growth rates by only 6%. Although C3 species grew 60% faster than CAM, the two groups showed no significant difference in their CO2 responses. High light increased average growth rates by 21%; high nutrients by 10%. The findings contrast with those noted by Granados and Korner (2002) and Wurth et al (1998a), who found significant positive responses of lianas to elevated CO2 and deep shade, opposed to high light intensities. Thus, epiphytes will pose a lower risk to forest turnover and carbon stock losses. CO2-water interactions CO2-water interactions have two sides: the CO2-driven stomatal response; and the interactions with weather; such as drought. Under elevated CO2 conditions, plants will always absorb more CO2 per unit of water lost; regardless of stomata respond. However, experimental evidence confirms stomata may not be as sensitive to CO2 as previously thought (Korner and Wurth, 1996; Lovelock et al, 1999). The increase in atmospheric CO2 over the last century has highlighted the dynamic relationships between CO2 gain and water loss. The evidence for this is within tree rings, in the form of stable carbon isotope signals. Hietz et al (2005) observed these changes in Amazonian trees, where a change in ∂3C over the past two centuries infers increased intrinsic water use efficiency. Traditionally, when water acts as a limiting factor, scientists have drawn upon an array of responses; such as stomatal closure; reduced photosynthesis and growth. However, it has been understood for decades that photosynthesis is less sensitive to reduced water potential than biomass growth. Most of the evidence is derived from non-woody plants (Korner, 2003a). Less water uptake reduces turgidity, which reduces tissue formation, eventually limiting CO2 uptake. Wurth et al (2005) completed an extensive inventory for 17 tropical tree species in both the dry and wet seasons in Panama. They found NSC pools to be largest when growth was lowest and smallest when growth reaches a maximum. This is counterintuitive to what is normally expected! It had been suggested that high NSC levels found in trees under growth limitations by environmental factors, such as drought, does not reflect source saturation by C, but a precaution strategy by which NSCs are stored in a reserve (Lewis et al, 2004a).

God’s Hands Behind Boxing Gloves Essay

It was after the 1960 Rome Olympics. A well-built man stood by a river and threw a gold medal into the waters. He had been rejected entrance in an all-white gathering, he who had brought pride to his nation.   Ã‚  Ã‚  Ã‚  Ã‚  Ã‚  Ã‚  Ã‚  Ã‚  Ã‚  Ã‚   The man was Muhammad Ali. He showed that no amount of worldwide recognition can be balm enough to the injustices and racism that was so prevalent during his time. The recognition that he earned at the Olympics was not ticket to equality, and he realized this instantly. He cared not for the medal, but the respect that his art and skill bring to the man, not the personality that the public knows. Muhammad Ali was not afraid to be seen with the much celebrated and controversial Malcolm X, and he fought his battles with as much confidence and gusto as he did within the ring and off it.   Ã‚  Ã‚  Ã‚  Ã‚  Ã‚  Ã‚  Ã‚  Ã‚  Ã‚  Ã‚   Dubbed as the â€Å"Athlete of the Century†, Muhammad Ali is known all over the world as one of the greatest boxers of all time. Indeed, with all bravado he dared call himself the Greatest. And he was, in many aspects not confined in the boxing ring.   Ã‚  Ã‚  Ã‚  Ã‚  Ã‚  Ã‚  Ã‚  Ã‚  Ã‚  Ã‚   Born as Cassius Clay Jr., he left behind his â€Å"slave name† and took up the name Muhammad Ali to assert his individuality and faith. His action has spurred black pride, as the man continued to prove to be more than just a boxer, but more importantly a black who is not afraid to succeed and fight for his rights. He refused to be inducted in the US Army as a stance against the war. He was imprisoned and stripped off his boxing title for his refusal, but he trained even behind bars and came back to recapture the championship and the people’s hearts. He staged his fights in developing countries that signaled a shift from first-world supremacy to an awareness of the existence of cities such as Manila and Kuala Lumpur. Muhammad Ali has visited and reached out to Afghanistan, North Korea, Cuba, Iraq and South Africa, going where there is strife and bringing goodwill.   Ã‚  Ã‚  Ã‚  Ã‚  Ã‚  Ã‚  Ã‚  Ã‚  Ã‚  Ã‚   Even after his popularity as a professional boxer has waned, his heart of gold shone brightly as ever. He promoted understanding at a time when there was discord in the streets, he embraced difference and stood his ground and pushed with his quiet way of battling racism – not through force but through tolerance. It was through not harboring hate that he gained peace of mind and freedom. He was a living example to all the blacks who wanted to be free. He has touched the lives, minds, and spirits of peoples from all over the world, from impoverished countries, to ghettoes, to aspiring boxers. He continues to make a difference in the lives of those around him, contributing to research centers and humanitarian works, gracing Make-A-Wish-Foundation and soup kitchens. Even if his claim to fame have been his powerful fists, more lasting was the kindness that his hands have shared to countless people. Muhammad Ali was known for wit, charm, and cocky, especially his famous proclamation, â€Å"I am the greatest†.   And the world agrees.

When To Use Accents and Diacritical Marks in Spanish

The most immediately obvious difference between written Spanish and written English is Spanishs use of written accents, and occasionally of diereses (also known as umlauts). Both of these features are known as diacritical marks. Beginning Spanish students usually learn right away that the main use of the accent is to help with pronunciation, and specifically in telling the speaker which syllable of a word should be stressed. However, accents also have other uses, such as distinguishing between certain homonyms, parts of speech, and indicating a question. The only use of the dieresis is to assist in pronunciation. Here are the basic rules for using the written accent and the dieresis: Stress The rules for determining which syllable should be stressed are quite simple in Spanish. Accents are used to indicate exceptions to the rules. Here are the basic rules: If a word ends in a vowel, the letter s, or the letter n, the stress is on the next to last syllable.In other words without an accent, the stress is on the last syllable. Simply put, if the stress is on a syllable other than that indicated above, an accent is used to indicate where the stress is placed. Following are a few examples, with the approximate pronunciation in phonetic English. Note that a vowel may either gain or lose an accent when a word is put into plural or singular form. See the rules on pluralization for other examples. examen (egg-SAH-men)exà ¡menes (eggs-SAH-men-ess)muà ±Ãƒ ³n (moon-YOHN)muà ±ones (moon-YOHN-ness)cancià ³n (kahn-SEEOHN)canciones (kahn-SEEOHN-ess) Distinguishing Homonyms Homonym pairs are separate words that have different meanings even though they sound alike. Here are some of the most common ones: de, of, from; dà © first- and third-person singular subjunctive form of dar, to give)el, the; à ©l, hemas, but; mà ¡s, moremi, my; mà ­, me;se, a reflexive and indirect object pronoun used in various ways; sà ©, I knowsi, if; sà ­, yessolo, only (adjective), single, alone; sà ³lo, only (adverb), solelyte, you (as an object); tà ©, teatu, your; tà º, you Demonstrative Pronouns Although the spelling reform of 2010 means they arent strictly necessary except to avoid confusion, accents also are traditionally used in Spanish on demonstrative pronouns to distinguish them from demonstrative adjectives. Talk about demonstrative parts of speech might sound like a mouthful, so its probably best to remember that in English were simply talking about the words this, that, these and those. In English, those words can be either adjectives or pronouns. In I like this book, this is an adjective; in I like this, this is a pronoun, since it stands for a noun. Here are the same sentences in Spanish: Me gusta este libro, I like this book. Me gusta à ©ste, translated as either I like this or I like this one. Note that when used as a pronoun, à ©ste traditionally has a written accent. In Spanish the demonstrative pronouns in the singular masculine form are à ©ste, à ©se, and aquà ©l, and the corresponding adjectives are este, ese, and aquel. Although distinguishing the meanings of these pronouns goes beyond the scope of this lesson, suffice it to say here that este/à ©ste corresponds roughly to this, while both ese/à ©se and aquel/aquà ©l can be translated as that. Items with which aquel/aquà ©l are used are farther from the speaker. Quiero aquel libro could be translated as I want the book thats over there. The following chart shows the various forms of the demonstrative pronouns (with the traditional accents) and adjectives, including the feminine and plural forms: Quiero este libro, I want this book. Quiero à ©ste, I want this one. Quiero estos libros, I want these books. Quiero à ©stos, I want these ones. Quiero esta camisa, I want this shirt. Quiero à ©sta, I want this one. Quiero estas camisas, I want these shirts. Quiero à ©stas, I want these ones.Quiero ese libro, I want that book. Quiero à ©se, I want that one. Quiero esos libros, I want those books. Quiero à ©sos, I want those ones. Quiero esa camisa, I want that shirt. Quiero à ©sa, I want that one. Quiero esas camisas, I want those shirts. Quiero à ©sas, I want those ones.Quiero aquel libro, I want that book over there. Quiero aquà ©l, I want that one over there. Quiero aquellos libros, I want those books over there. Quiero aquà ©llos, I want those ones over there. Quiero aquellas camisas, I want those shirts over there. Quiero aquà ©llas, I want those ones over there. There are also neuter variations of these pronouns (eso, esto, and aquello), and they are not accented because there are no corresponding neuter adjective forms. Interrogatives: A number of words are accented when they are used in a question (including an indirect question) or exclamation, but they arent otherwise accented. Such words are listed below:  ¿Adà ³nde? Where (to)? ¿Adà ³nde vas? Where are you going? ¿Cà ³mo? How? ¿Cà ³mo està ¡s? How are you? ¿Cuà ¡l?  ¿Cuà ¡les? Which one? Which ones? ¿Cuà ¡l es mà ¡s caro? Which one is more expensive? ¿Cuà ¡ndo? When?  ¿Cuà ¡ndo sales? When are you leaving? ¿Cuà ¡nto?  ¿Cuà ¡nta?  ¿Cuà ¡ntos?  ¿Cuà ¡ntas? How much? How many?  ¿Cuà ¡ntos pesos cuesta el libro? How many pesos does the book cost? ¿Dà ³nde? Where?  ¿De dà ³nde es usted? Where are you from? ¿Por quà ©? Why?  ¿Por quà © vas? Why are you going? ¿Quà ©? What? Which?  ¿Quà © libro prefieres? What book do you prefer? ¿Quià ©n?  ¿Quienes? Who? Whom?  ¿Quià ©nes quieren mi libro? Who wants my book? Diereses: The dieresis (or umlaut) is used above the u when the u is sounded in the combinations of gà ¼i or gà ¼e. Without the umlaut, known as la dià ©resis or la crema in Spanish, the u would be silent, serving only to indicate that the g is pronounced as a hard g rather than similar to the j. (For example, guey with no umlaut would sound something like gay.) Among the words with umlauts are vergà ¼enza, shame; cigà ¼eà ±a, stork or crank; pingà ¼ino, penguin; and agà ¼ero, prediction.

Germany as a Victim - 1451 Words

Germany as a Victim On the 16th June the German government, lead by Count Brockdorff-Rantzau, were presented with the Treaty of Versailles. They were originally given 14, and then 21 days, to agree to it. The treaty, which included some 440 Articles, was not as vindictive as Clemenceau had wanted nor as moderate as Lloyd George would have wished. It certainly fell far short of the conciliatory features of Wilsons fourteen point proposals. (Evans and Jenkins) However, in order to decipher whether the end results were destructive to Germany or not, it is necessary to asses not only the main points of the treaty (including military provisions, territory, financial provisions, war guilt and the†¦show more content†¦The most obvious term was the return of Alsace-Lorraine to France. Schleswig was also returned to Denmark after it had been annexed by Bismark. Also Eupen and Malmedy were to go to Belgium. This act is thought to be justified by the fact that much of Belgiums industry was destroyed du ring to war, whilst Germany remained relatively untouched. The Saar Basin was also to be overseen by the League of Nations, the Saar coalfield by France, for fifteen years. This amounted to 13% of German territory and 12% of the population - half of which were ethnic Germans. Lloyd George claimed that I cannot conceive any greater cause of future war than that the German people should be surrounded by small states . . . each of them containing large masses of Germans clamouring for reunion with their homeland It also took away much of their industry including 48% iron ore, 16% coal, and15% agricultural production. 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