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Carbon allocation patterns in plants and plant ecosystems Walton, Anne Barber 1995

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C A R B O N A L L O C A T I O N PATTERNS IN PLANTS A N D P L A N T ECOSYSTEMS by A N N E BARBER W A L T O N B.Sc. Honours, Queen's University, 1991 A THESIS SUBMITTED IN PARTIAL F U L F I L L M E N T OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in THE F A C U L T Y OF G R A D U A T E STUDIES (Department of Botany) We accept th|s thesi^as conforming to the required standard THE UNIVERSITY OF BRITISH C O L U M B I A July 1995 © Anne Barber Walton, 1995 In presenting this thesis in partial fulfilment of the requirements for an advanced degree at the University of British Columbia, I agree that the Library shall make it freely available for reference and study. I further agree that permission for extensive copying of this thesis i o r scholarly purposes may be granted by the head of my department or by his or her representatives. It is understood that copying or publication of this thesis for financial gain shall not be allowed without my written permission. Department of RftTAOy The University of British Columbia Vancouver, Canada Date hjuuTAJudt I. I 9 9 3 0 1 DE-6 (2/88) A B S T R A C T A study of plant carbon allocation patterns at the ecosystem and cellular levels revealed interactions between ecological and physiological processes. At the global level, ecosystem net' primary production was allocated to lignin and holocellulose synthesis in plant life-forms and plant organs. Forty-eight percent of the global net primary production (carbon) was estimated to be associated with the synthesis of lignin and holocellulose, which are the primary constituents of plant cell walls. The primary ecosystems associated with this lignin and holocellulose production were tropical forests, Northern Hemisphere forests, and savannas. Associated with the annual net production of lignin, it was estimated that l x l O 1 4 moles of nitrogen are reassimilated through the prephenate-arogenate junction which links primary carbon metabolism to secondary carbon metabolism. Estimates of annual net primary production associated with plant life-forms and plant organs indicated that the two greatest contributors were herbaceous and nonvascular aquatic (non-macrophyte) plants. Estimates may provide a basis for future models of pools and fluxes of carbon in the global carbon cycle. At the cellular level, treatment of parsley (Petroselinum crispum L.) cell cultures with the Phytophthora megasperma elicitor caused an immediate increase in the rate of respiratory carbon dioxide evolution in the dark that corresponded to the activation of phosphofructokinase and glucose-6-phosphate dehydrogenase, the key enzymes in the regulation of carbohydrate flow to glycolysis and the oxidative pentose phosphate pathway, respectively. The increased rate of carbon dioxide evolution and the activation of phosphofructokinase and glucose-6-phosphate dehydrogenase are maintained for the duration of the experiments indicating long-term stimulation of respiration through both glycolysis and the oxidative pentose phosphate pathway. A 23% decrease in the C6 :C i ratio 60 minutes after elicitation was consistent with increased contribution of the oxidative pentose phosphate pathway to cellular respiration. Long-term activation of the oxidative pentose phosphate pathway following elicitation could serve to maintain the pools of substrates necessary during activation of the shikimic acid pathway, leading to the production of defensive compounds. T A B L E OF CONTENTS 111 A B S T R A C T i i T A B L E OF CONTENTS i i i LIST OF T A B L E S viii LIST OF FIGURES ix ABBREVIATIONS x A C K N O W L E D G M E N T S xi F O R W A R D xii IN L O V I N G M E M O R Y xiii CHAPTER 1. Introduction 1 Plant cell carbon allocation 1 Whole plant carbon allocation 4 Plant ecosystem carbon allocation 5 CHAPTER 2. Annual Net Flow of Carbon to Lignin and Holocellulose Synthesis 7 Introduction 7 Methods 9 Step 1: Estimation of Global Plant Net Primary Production Associated with Ecosystems 9 Step 2: Estimation of Ecosystem Net Primary Production Associated with Plant iv Life-Forms and Plant Organs 9 Forest 15 Tropical Humid 15 Tropical Seasonal 20 Mangal 22 Temperate Evergreen/Coniferous 22 Temperate Deciduous/Mixed 24 Boreal Coniferous (closed) 25 Boreal Coniferous (open) 26 Plantation 27 Plantation (<10 y) 28 Plantation (11 y and older) 29 Temperate Woodland 30 Chaparral, Maquis and Brushland 31 Savanna 32 Nonseasonal 33 Seasonal 33 Hyperseasonal 34 Temperate Grassland 35 Tundra 37 Polar Desert 37 High Arctic/Alpine 38 Low Arctic/Alpine 38 Desert 39 Desert and Semidesert Scrub 39 Extreme (sandy, hot and dry) 40 Extreme (sandy, cold and dry) 40 Lake and Stream 41 Swamp, Marsh, and Estuary 42 V Freshwater Swamp 42 Freshwater Marsh 44 Saltwater Marsh/Estuary 44 Algal Bed and Reef 45 Peatland 45 Bog 46 Fen 46 Cultivated Land 47 Herbaceous Reproductive Crops 47 Herbaceous Shoot Crops 48 Herbaceous Root/Tuber Crops 48 Woody Reproductive Crops 49 Woody Leaf Crops 50 Human Area 50 Marine 51 Step 3: Estimation of Plant Organ Net Primary Production Associated with Lignin and Holocellulose 52 Step 4: Estimation of Lignin and Holocellulose Net Primary Production Associated with Carbon 52 Results and Discussion 56 Global production of lignin and holocellulose 56 Ecosystem contribution to global net flow of carbon to lignocellulose 56 Lignocellulose production in the forest ecosystem 58 Lignocellulose production in the savanna ecosystem 64 Lignocellulose production in the cultivated land ecosystem 66 Lignocellulose production in the grassland ecosystem 66 Lignocellulose production in the swamp, marsh, and estuary ecosystem 67 Lignocellulose production in the marine ecosystem 68 Lignin production and the global nitrogen cycle 68 Allocation of global net primary production 70 Plant life-forms 70 Angiosperm woody plants 72 Gymnosperm woody plants 72 Herbaceous plants 73 Macrophytic plants 73 Nonvascular plants 74 Plant organs 74 Ratios of belowground net primary production to aboveground net primary production 76 Implications for modeling global plant net primary production 79 Estimate caveats 81 First caveat 81 Second caveat 82 Third caveat 82 Fourth caveat 82 CHAPTER 3. Pathogen-Induced Alterations of Respiratory Carbon Flow 84 Introduction 84 Methods 86 Cell Culture and Elicitor 86 Gas Exchange Experiments 86 Metabolite Experiments 87 C6:C] Ratios: Analysis of 1 4 C 0 2 Evolution 87 Other Methods 88 Results 89 Respiratory CO2 Evolution 89 Metabolites 89 C6:Ci Ratios of 1 4 C 0 2 Evolution 93 Vll Discussion 95 Immediate Events Following Elicitor Treatment 95 Long-Term Events Following Elicitor Treatment 96 CHAPTER 4. Conclusions 99 Global carbon allocation Lignocellulose production 99 Production of plant life-forms and plant organs 100 Implications for models 100 Pathogen-induced changes in plant carbon allocation 101 B I B L I O G R A P H Y 102 APPENDIX. Global Net Primary Production Partitioning (Microsoft Excel Spreadsheet) 122 viii LIST OF TABLES Table 2.1 Surface area and annual net primary production values of global ecosystems 11 Table 2.2 Global classification of plant life-forms and plant organs 13 Table 2.3 Percentage of annual ecosystem net primary production (NPP, g dry matternr2) estimated to be associated with plant life-forms and plant organs: Belowground-Aboveground Method 16 Table 2.4 Percentage of annual ecosystem net primary production (NPP, g dry matternr2) estimated to be associated with plant life-forms and plant organs: Plant Life-Form Method 36 Table 2.5 Relative amounts of holocellulose and lignin in terrestrial plant organs 53 Table 2.6 Annual net flow of photosynthetically fixed carbon to global lignin and holocellulose synthesis 57 Table 2.7 Lignocellulose content, rate of C turnover, and C-sink potential of plant organs 59 Table 2.8 Estimated ecosystem ratios of belowground net primary production (BNPP) to aboveground net primary production (ANPP) 77 Table 3.1 Effect of Pmg elicitor (30 i^g-mL"1) on the Cg:Ci ratios of 1 4 C 0 2 evolution from cell suspensions of Petroselinum crispum. (mean ± SD) 94 IX LIST OF FIGURES Figure 1.1 Diagrammatic representation of the respiratory pathways participating in the biosynthesis of phenylpropanoid compounds 3 Figure 2.1 Four-step process to estimate the flow of net annual photosynthetic carbon fixation to global lignin and holocellulose synthesis in plants 10 Figure 2.2 Plant organ contribution to lignin production in forest (A), savanna (B), cultivated land (C), grassland (D), and swamp, marsh and estuary (E) ecosystems 60 Figure 2.3 Plant organ contribution to holocellulose production in forest (A), savanna (B), cultivated land (C), grassland (D), and swamp, marsh and estuary (E) ecosystems 61 Figure 2.4 Ecosystem contribution to annual net primary production of angiosperm woody (A), gymnosperm woody (B), herbaceous (C), and nonvascular aquatic (non-macrophyte) (D) plant dry matter (DM) 62 Figure 2.5 Diagrammatic representation of nitrogen reassimilation during lignin biosynthesis at the prephenate-arogenate junction. 69 Figure 2.6 Annual net primary production (NPP) associated with plant life-forms and plant organs 71 Figure 3.1 Representative traces of the changes in the rate of CO2 evolution in cell suspensions of Petroselinum crispum treated with water (A) or Pmg elicitor 30 (igrnL- 1) (B) 90 Figure 3.2 The intracellular levels of Fru-6-P (A) and Fru-l,6-bisP (B) and the ratio of Fru-l,6-bisP:Fru-6-P (C) in cell suspensions of Petroselinum crispum treated with Pmg elicitor (30 pLgmL'1) (O) and water (•). 91 Figure 3.3 The intracellular levels of Glc-6-P (A) and 6-PG (B) and the ratio of 6-PG:Glc-6-P (C) in cell suspensions of Petroselinum crispum treated with Pmg elicitor(30 pig-mL-1) (O) and water (•). 92 A B B R E V I A T I O N S A N P P , aboveground net primary production B N P P , belowground net primary production C , carbon CO2, carbon dioxide D A H P , 3-deoxy-D-ara£mo-heptulosonate-7-phosphate ddH.20, distilled deionized water D M , dry matter E4P, erythrose-4-phosphate Fru-6-P; fructose-6-phosphate Fru-1,6-bisP, fructose-1,6-bisphosphate G 6 P D H , glucose-6-phosphate dehydrogenase (EC 1.1.1.49) Glc-6-P, glucose-6-phosphate holocellulose, cellulose plus hemicellulose I R G A , infrared gas analyzer lignocellulose, lignin plus cellulose N , nitrogen N P P , net primary production OPP, oxidative pentose phosphate Pg, petagram, 1 0 1 5 g Pmol, petamole, 10 1 5 mol 6-PG, 6-phosphogluconate P F K , phosphofructokinase (EC 2.7.1.11) Pmg, Phytophthora megasperma elicitor T C A , tricarboxylic acid T E M , terrestrial ecosystem model A C K N O W L E D G M E N T S I have had the opportunity to befriend many people during my four-year stay in Vancouver. Of these people, a few have enhanced my life personally and academically thus helping me to attain the long-awaited completion of my Master of Science degree in Botany. Dr. David H . Turpin, my supervisor, offered me the opportunity of a lifetime to become a part of the Botany Department of the University of British Columbia. Through his initial supervision and ideas, he opened my eyes to the great wonders of the plant kingdom. His financial support, through the National Science and Engineering Research Council of Canada, allowed me to continue my degree until December of 1994. Dr. Eric G. Norman was the necessary catalyst who helped me to re-assess the methodologies for both the global and physiological projects. The collaboration with Dr. Norman was the highlight of my program and left me with the idea that almost anything is possible if one is open to looking for alternate problem-solving methods. Drs. Bruce A . Bohm and Brian E. Ellis provided me with perhaps the most important gift, that of personal support and belief in my abilities to expand and complete the global research. Dr. Carl J. Douglas was kind enough to teach a neophyte the techniques of higher plant cell culture and provide the Pmg elicitor. Heather Huppe, David Gauthier, Troy Sitland, and Jody Holmes, among other members of the Turpin laboratory, were a joy to work with on a daily basis and were my scientific mentors in many ways. The members of the Botany office, including Dr. Iain E. P. Taylor, the secretarial staff, and especially R. Patrick Harrison, have provided the much needed personal support to deal with the many deadlines accompanying a Master of Science degree. The Botany Department has generously provided financial support since January of 1995. The University of British Columbia librarians, especially Jim Harris, have helped me to obtain references from obscure publications and to maintain a semi-reasonable fines record while consistently having between 100 and 200 books checked out of the library. There are a few personal friends who have given me tremendous support in completing my thesis: Rob Knobel, Les Shewchuk, Darren Goetze, Jens Haeusser, and Alan Reid. Finally, my family has maintained the loving support of my capabilities to finish this degree and return to the U.S.A. Xll FORWARD The research presented in this thesis is the result of a collaborative effort with Dr. Eric G. Norman who joined the laboratory group of Dr. David H. Turpin in January of 1993. Before this collaboration began, I had written a preliminary paper which was the starting point for the research presented in Chapter 2. I had also conducted preliminary gas exchange experiments using Petroselinum crispum cell cultures and the Phytophthora megasperma elicitor which indicated elicitor-induced increases in CO2 evolution. The results presented in Chapter 2 are based upon methodologies which were largely developed by me. With the exception of the relation between net primary production and percent tree cover in the Human Area ecosystem (p. 50), I conducted all research presented in this chapter, developed the methodology presented in this chapter. The literature search, data collection, analysis, and presentation, and writing were carried out by me. The text, figures, and table of Chapter 3 are taken directly, with the exception of formatting, from the article published in the journal, Plant Physiology (106): 1541-1546, 1994. Permission to do this was granted from both the publisher of Plant Physiology, The American Society of Plant Physiologists, and the first author of the article, Dr. Eric G. Norman ~ A „. . The writing for the first draft of the manuscript was shared equally between Dr. Norman and me. I was responsible for parts of the introduction, the methods and results sections pertaining to Figs. 3.1, 3.2, and 3.3, and the sections of the discussion dealing with short-term events following elicitation. The literature search, data collection, analysis, and presentation of Figs. 1.1, 3.1, 3.2, 3.3, and data not shown sections dealing with metabolites were carried out by me. Dr. Norman conducted the research presented in Table 3.1, the data not shown dealing with pH changes following elicitation, and the data not shown dealing with [U- 1 4 C] phenylalanine and [6- 1 4C] glucose. Dr. Norman was responsible for the majority of the revisions to the text. xiii IN L O V I N G M E M O R Y I present the work in this thesis in loving memory of my grandfather, William Dickson Macdonald, who passed away in Gainesville, Florida on April 15, 1994. In many ways, my academic success is attributed to him. Throughout my childhood, he treated me as an equal. I clearly remember his faith in my abilities when, in my pre-teenage years, he allowed me to record the exam marks of his law students in his record book. As I progressed through middle school and then the International Baccalaureate Program in high school, he supported each academic project I accomplished. Though he was born, raised, and educated in Canada, surprisingly, he was the only family member to object to me leaving the U.S.A. to begin undergraduate studies in Canada. In August of 1993, however, I recall earning his approval when I showed him the physiological work which I had presented at a scientific meeting. He smiled and repeated an idea from my poster which contained the phrase, "phenylpropanoid metabolism," though slightly mispronounced. One day, perhaps I will walk in the footsteps of my grandfather and pass on his inspiration to my future children and grandchildren. CHAPTER 1. INTRODUCTION Plant carbon (C) allocation may be considered in levels, ranging from the small scale (cellular) to the large scale (global). At each level, regulatory mechanisms maintain the balance of C allocation between sources and sinks as the plant grows and encounters a variety of changing conditions, loosely termed "stresses." These stresses may be abiotic or biotic. Abiotic stresses arise from changes in climate or the levels of pollutants and inorganic nutrients. Biotic stresses arise from the interactions between plants and animals, insects, fungi, bacteria, or other plants. Changes in C allocation patterns at one level are usually reflected in neighboring levels. Therefore, by studying the mechanisms underlying the regulation of allocation within and among levels, we may predict more accurately how and when changes may occur in the future. The following is a general review of plant C allocation patterns that occur at each level with implications for understanding interactions between levels. Plant Cell Carbon Allocation Carbon allocation among constituents in a plant cell is dependent foremost upon the primary functions of the given cell. A photosynthesizing leaf cell of a C3 plant, for example, cycles between accumulation of C in the form of storage product (starch) during the day and oxidation to sucrose at night. A root cell has greater C stores during the winter which are hydrolyzed to sucrose for export to shoot growth at the beginning of the growth season (Hocking 1989). These two examples illustrate the cyclical nature of C allocation in whole cells which occurs daily (former example) or seasonally (latter example). Another feature of these examples is in the allocation of C to different subcellular compartments. In these examples, C is stored as starch in plastids but may be utilized in other compartments, including the cytoplasm, mitochondria, and other areas. The process of C utilization involves a tightly regulated set of biochemical reactions which compete for these C stores. One of the most complex pathways that is vital to the production of cellular energy and C metabolites is that of respiratory metabolism. Carbon may be allocated through respiratory metabolism via one of two routes, namely glycolysis and the oxidative pentose phosphate (OPP) pathway. Regulation of C flow through these two pathways is dependent upon the cellular demands for products. The primary end-products of glycolysis are 3-C organic compounds, ATP, and NAD(H). Mitochondrial respiration places a considerable demand on the plant cell for these substrates. The OPP pathway may provide the same products as glycolysis, since the pathways share common intermediates, in addition to 4-C, 5-C, 6-C, and 7-C compounds and NADP(H) . These products are used in the anabolic reactions leading to the synthesis of nucleotides and aromatic amino acids, among other compounds. Since the two respiratory pathways may yield products which are utilized for quite different functions, regulation of C allocation between them should be tightly linked to the given cellular demands. Environmental stresses place different demands upon plants that may lead to shifts in the C allocation patterns between glycolysis and the OPP pathway. Secondary plant compounds, for example, are synthesized from pathways which utilize different respiratory products. Terpene biosynthesis requires acetate which is derived from either glycolysis or the OPP pathway. Phenylpropanoid metabolism, which leads to the biosynthesis of such compounds as lignin and flavonoids, requires phosphoeno/pyruvate (PEP), derived from either glycolysis or the OPP pathway, and erythrose-4-phosphate (E4P), derived from the OPP pathway (Fig. 1.1). A review of the literature revealed evidence supporting the idea of differential C allocation among respiratory pathways in response to environmental stress. Under long-term CO2 enrichment of Picea abies, activity of glucose-6-phosphate dehydrogenase, a regulatory enzyme of the OPP pathway, declined while activities associated with tricarboxylic acid (TCA) cycle enzymes (fumarase and NAD-malic enzyme) increased (Van Oosten et al. 1992). These results are consistent with decreased C flow through the OPP pathway but increased C flow through glycolysis. Increased glycolytic C flow was also indicated in Arabidopsis (Yang et al. 1993). Increased transcription of mRNA coding for the glycolytic enzyme glyceraldehyde-3-phosphate dehydrogenase was demonstrated under two stress treatments, namely heat-shock and anaerobiosis. Thus, a general C allocation response in plants confronted with new stresses may be to alter C allocation between glycolysis and the OPP pathway. This has implications for c r c o 2 2 Glc-6-P I I 1 Fru-6-P*#-G | PFK c 1 Fru-1,6-bisP 0 1 y s i s J PK 2 PEP I I 2 PYR I / TCA Cycle j OPP Pathway 2 NADPH 2 6-PG G6PDH t 6-PGDH | 2 NADPH + 2 C r C 0 2 2 R5P 1 Fru-6-P + 1 E4P 1 E4P 1 PEP 1 DAHP synthase 1 DAHP 1 PEP ^— 1 NADPH T ' 1 PHE Shikimic Acid Pathway PHE NADPH 1 FC Phenylpropanoid Metabolism Figure 1.1 Diagrammatic representation of the respiratory pathways participating in the biosynthesis of phenylpropanoid compounds. (Abbreviations: DAHP, 3-deoxy-D- arabmo-heptulosonate-7-phosphate; E4P, erythrose-4-phoshate; FC, furanocoumarins; G6PDH, glucose-6-phosphate dehydrogenase; PFK, phosphofructokinase; 6-PG, 6-phosphogluconate; 6-PGDH, 6-phosphogluconate dehydrogenase; PHE, phenylalanine; PK, pyruvate kinase; PYR, pyruvate; R5P, ribulose-5-phosphate; TCA, tricarboxylic acid). whole plant C allocation patterns since increased allocation to respiratory C O 2 efflux may lead to decreased allocation to tissue and organ sinks. Whole Plant Carbon Allocation Whole plant C allocation patterns are a function of the relative demands associated with the various plant organ C sinks. Patterns of C allocation to organ sinks are dynamic and vary with plant life-form (Mooney 1972; Pitelka 1977), plant age (Chalmers and van den Ende 1975; Singh et al. 1993) season (Chung and Barnes 1980; Schier 1970), stage of reproductive maturation (Bazzaz et al. 1987; Marcelis 1992), altitudinal location (Korner and Renhardt 1987), nutritional status (Ingestad and Agren 1991; Nadelhoffer et al. 1985; Waring et al. 1985), and light availability (Pasumarty and Fountain 1993; Waring et al. 1985). Whole plant C allocation processes are therefore difficult to model and predict since they may be altered in response to combinations of changing environmental and growth conditions. However, whole plant C allocation has been classified into three types based up the function of the given sink: growth, defense, and reproduction (Clark and Harvell 1992). Carbon allocation changes associated with growth, defense, and reproduction have been documented in various studies of plant stress. Changes in C allocation related to plant growth included increased allocation to belowground plant organs in a higher C O 2 climate (Norby et al. 1992) and higher ozone environment (Qiu et al. 1992), increased growth in elevated C O 2 (Bazzaz et al. 1990; Norby et al. 1992), and decreased overall growth related to ozone exposure (Karnosky et al. 1992; Qiu et al. 1992). Changes in allocation related to plant defense included increased production of secondary plant compounds (Lindroth et al. 1993) and decreased growth (Paul and Ayres 1986). Finally, changes related to reproduction included increased reproductive output associated with a number of stresses (Smit 1980, cited in Grime and Campbell 1991). Overall, it appears that whole plant C allocation is sensitive to a number of environmental stresses. This may have implications for C allocation patterns at the ecosystem level since whole plant C alterations may be advantageous for some life-forms while being a hindrance for others. The outcome may be changes in population structure and cycling of C within ecosystems. Plant Ecosystem Carbon Allocation Estimates of ecosystem C allocation are a function of the C allocated among all plant components within a given ecosystem. Ecosystem C allocation is therefore a difficult process to study since life-forms may include woody, herbaceous, and nonvascular plants adapted of many specialized growth-forms (Whittaker 1970). One of the prime goals of the International Biological Program, established in 1964, was to improve our understanding of biological productivity in global ecosystems, both aquatic and terra firma. At the end of the program, in 1974, a series of books was published. Carbon allocation patterns related to biomass, litterfall, and primary production were presented for the main ecosystems. An obvious missing piece of information was that of primary production allocated to belowground organs. This was amplified in the global estimate of C allocation among terrestrial ecosystems by Ajtay et al. (1979) which utilized information collected during the ten years of study for the International Biological Program series. In the global estimate, Ajtay et al. (1979) acknowledged that belowground information was lacking and made an undocumented estimate of its contribution to the global C budget. Current field research is focused upon improving the estimates of C allocation within ecosystems by studying the role of fine roots (Nadelhoffer and Raich 1992) and herbivory (Pastor and Naiman 1992) in the C budget. Modeling research is focused upon estimating ranges of carbon allocation patterns based upon multiple biotic and abiotic parameters (Raich etal. 1991). Predictions of changes to C pools associated with different plant life-forms will improve with greater understanding of C cycling processes in natural ecosystems. Small-scale studies on tundra and marsh (Mooney et al. 1991) and tropical forest ecosystems (Korner and Arnone III 1992) were conducted to estimate potential changes to C allocation in artificially elevated CO2 environments. In both marsh (C3-plants) and tropical forest systems, C allocation to belowground organs was increased whereas overall long-term stimulation of growth was experienced only in the marsh (C3) ecosystem. Since the rate of climate change is unlikely to be as great as that imposed on plant systems in controlled studies, current research suggests that plant processes may be adaptable to the graduated changes in climate which are predicted to occur over the next 100 years. In the following two chapters, I present information that will fill gaps in our knowledge at two levels of plant C allocation, the ecosystem and the cellular. At the ecosystem level, I present a first estimate of the annual plant C allocation to lignin and holocellulose synthesis. Lignin and holocellulose are the two most abundant plant polymers and as such comprise the bulk of cell wall material. Cell wall degradation is the limiting factor to C turnover in plant litter; therefore, this estimate will provide the information required to understand current patterns of C turnover times from global detrital pools. At the cellular level, I present the first evidence for immediate changes in C allocation that accompany a representative plant-pathogen interaction. Overall, this thesis presents research which elucidates patterns of C allocation to growth and reproduction on the global level and to defense at the cellular level. CHAPTER 2. A N N U A L NET FLOW OF C A R B O N TO LIGNIN A N D H O L O C E L L U L O S E SYNTHESIS INTRODUCTION The human population is increasing at an exponential rate and is expected to nearly triple from its current level of 5.5 billion by the end of the 21 s t century (Schlesinger 1991). Anthropogenic activities associated with this growth will have serious consequences for global biodiversity and climate change as toxic wastes accumulate and global biogeochemical cycles of C and N are altered. Already there are signs of widespread environmental destruction as evidenced by the declining global populations of amphibians. Large-scale pollution events such as toxic waste accumulation in landfills, oil spills, and tropospheric smog, receive a great deal of public attention. However, the cumulative effects of smaller-scale, individual pollution events may be equally potent and therefore receive much attention from the scientific community. One of the most widely studied pollutants of this kind is CO2, which has been linked to global warming. The atmospheric concentration of CO2, currently at 355 ppm (Emanuel et al. 1994), is increasing at a rate of 0.4% per year (Schlesinger 1991) due to the combined net biospheric release of CO2 from fossil fuel combustion, estimated at 5 petagrams (Pg=1015g) C y 1 (Rotty and Masters 1985), and destruction of vegetation from land-use change, primarily through tropical deforestation, estimated to be releasing 1.8 Pg C - y 1 (Houghton et al. 1987; Brown et al. 1993). Associated with the increased levels of greenhouse gases are the predicted increases in the mean annual temperature (2.8-5.2 °C) and mean annual precipitation (7.1-15.8%) expected in a 2xC02 environment. This environment may occur by 2075 if the current rates of CO2 release to the atmosphere remain steady (Shafer and Schoeneberger 1991). Recent studies of plant growth in response to C02-enriched environments suggest an initial positive response of terrestrial plant growth to increasing CO2 concentrations (Bazzaz 1990; Hocking and Meyer 1991; Pastor and Post 1988). To predict long-term responses of global ecosystems under a 2xC02 environment, models are more frequently being employed. Many of the current C cycling models are based upon plant net primary production (NPP) studies from the 1970s, and it is likely that the methods employed may have significantly underestimated contribution from belowground organs (Nadelhoffer and Raich 1992). Another piece of information missing from the older ecological studies is that of total NPP associated with the many types of plant life-forms and plant organs. Plant organs contain differential amounts of the structural C polymers, lignin and holocellulose; thus, this information may be crucial to modeling functions of the C cycle since the quality of C in plant organ litterfall dictates the residence times of C in pools of organic matter (Melillo et al. 1982). Global location of plant litter in the unique climates of different ecosystems also plays a role in regulating C residence times. Turnover time and decomposition rates of plant litter in tropical climates, for example, are faster than in temperate climates (Cebrian and Duarte 1995; Schimel et al. 1994). In this chapter, I present an estimate of the annual net flow of C to global lignin and holocellulose synthesis in plant life-forms and plant organs. Potential global C sinks of lignin and holocellulose in different plant organs are inferred from the global location of plant organ production and the relative rates of organ decomposition. Results are also discussed with reference to current research that highlights the impact of predicted changes in the areal extent of global ecosystems upon the total production of plant life-forms. METHODS Information collected during a comprehensive literature review was used to estimate the flow of the annual net photosynthetic carbon fixation to global lignin and holocellulose (cellulose plus hemicellulose) synthesis. Figure 2.1 outlines the four steps involved in these processes. Step 1: Estimation of Global Plant Net Primary Production Associated with Ecosystems The first step involved estimation of the annual NPP of plant dry matter (DM) associated with global ecosystems. The ecosystem classifications and corresponding areal and NPP values of Ajtay et al. (1979) served as a basis for estimating the allocation of plant NPP (DM) among the global ecosystems (Table 2.1). Step 2: Estimation of Ecosystem Net Primary Production Associated with Plant Life-Forms and Plant Organs A database was creatd from 160 ecological studies, and references representing the 37 ecosystems and used to estimate the proportion of the ecosystem NPP (as classified in Table 2.1) associated with the various plant life-forms and organs listed in Table 2.2. Two methods were used to apply NPP information from the published studies and references to the plant life-forms and organs in each ecosystem, the Belowground-Aboveground method and the Plant Life-Form method. The Belowground-Aboveground method was used if the plant NPP information was presented separately for aboveground and belowground plant organs. For each ecosystem, the percentage of the NPP associated with aboveground (ANPP) and belowground plant life-forms (BNPP) was estimated using the BNPP:ANPP ratios obtained from published studies which included at least the dominant plant life-forms. Estimates of ANPP associated with the various plant life-forms and organs (%ANPP) were obtained from ecological studies and 10 Global Annual NPP (DM) STEP1 Ecosystem Annual NPP (DM) STEP 2 Plant and Organ Annual NPP (DM) STEP 3 Lignin and Holocellulose Annual NPP (DM) STEP 4 Lignin and Holocellulose Annual NPP (C) Figure 2.1 Four-step process to estimate the flow of net annual photosynthetic carbon fixation to global lignin and holocellulose synthesis in plants. (C, carbon; D M , dry matter; NPP, net primary production) Table 2.1 Surface area and annual net primary production values of global ecosystems. [* indicates ecosystem modifications from Ajtay et al. (1979) as noted in methods; A , angiosperm, G, gymnosperm; Aalgal bed and reef, marine, and estuary values estimated by Whittaker and Likens, 1973, 1975 and Whittaker 1975, cited in Ajtay etal. (1979)] S U R F A C E T O T A L A R E A NPP E C O S Y S T E M ECOSYSTEM-TYPE (xl0 1 2 m 2 ) (x l0 1 5 g DM) Forest Tropical humid 10 23 Tropical seasonal 4.5 7.2 Mangal 0.3 0.3 Temperate evergreen/coniferous 3 4.5 Temperate deciduous/mixed 3 3.9 Boreal coniferous (closed) 6.5 5.53 Boreal coniferous (open) 2.5 1.63 Plantation <10 y* 0.3 0.53 Plantation 11 y and older-A* 0.2 0.41 Plantation 11 y and older-G* 1.0 1.68 Total 31.3 48.68 Temperate woodland 2 3 Chaparral, maquis, brushland 2.5 2 Savanna* Nonseasonal 6 12.6 Seasonal 10.5 12.95 Hyperseasonal 6 13.8 Total 22.5 39.35 Temperate Grassland Moist 5 6 Dry 7.5 3.75 Total 12.5 9.75 Tundra Polar desert 1.5 0.04 High arctic/alpine 3.6 0.54 Low arctic/alpine 4.4 1.54 Total 9.5 2.12 Desert Desert and Semidesert 21 3.0 Extreme (sandy, hot and dry) 8 0.08 Extreme (sandy, cold and dry) 1 0.05 Total 30 3.13 Lake and stream 2 0.8 Swamp, marsh and estuary* Freshwater swamp 0.3 0.89 Freshwater marsh 0.2 0.43 Saltwater marsh/estuaryA 2.9 8.03 Total 3.4 9.35 Algal bed and reefA 0.6 1.6 Table 2.1 Surface area and annual net primary production values of global ecosystems. (Continued) S U R F A C E T O T A L A R E A NPP E C O S Y S T E M ECOSYSTEM-TYPE (x l0 1 2 m 2 ) ( x l 0 1 5 g DM) Peatland* Bog 1.4 1.39 Fen 0.1 0.11 Total 1.5 1.5 Cultivated Land* Herbaceous reproductive crops 13.5 12.67 Herbaceous shoot crops 0.5 0.51 Herbaceous root/tuber crops 0.9 0.84 Woody reproductive crops 1.0 0.92 Woody leaf crops 0.1 0.11 Total 16 15.05 Human Area 2 0.4 MarineA 508.0 51.3 G L O B A L TOTAL 659.3 188.03 13 Table 2.2 Global classification of plant life-forms and plant organs. [* petiole of submergent/floating-leaved macrophyte similar in structure to submergent stalks (Riemer 1984) therefore included with stalk organs; A , angiosperm; G, gymnosperm] P L A N T LIFE- PLANT LIFE-FORM P L A N T F O R M DESCRIPTION O R G A N Vascular-A Woody Trees Reproductive Shrubs Leaf (Terrestrial and Epiphytic) Stem Branch Fine root Coarse root Herbaceous Graminoids, Reproductive Forbs, Leaf Pteridophytes Stalk (Terrestrial and Epiphytic) Fine root Coarseroot/ rhizome/tuber Emergent Macrophyte Graminoids Reproductive Forbs Leaf Pteridophytes Stalk Fine root Coarse root/ rhizome S ubmergent/Floating- Graminoids Reproductive Leaved Macrophyte Forbs Leaf Pteridophytes Petiole/Stalk* Fine root Coarse root/ rhizome Vascular-G Woody Trees, Shrubs (Terrestrial) Reproductive Leaf Stem Branch Fine root Coarse root Nonvascular Aquatic-Macrophyte Macroalgae Aboveground Aquatic-Non- Phytoplankton, periphyton Aboveground macrophyte Terrestrial Cryptogams Aboveground references as described for the given ecosystem. Estimates of percent BNPP (%BNPP) associated with non-woody vascular plant life-forms were obtained by multiplying the B N P P : A N P P ratios for herbaceous and/or macrophytic plants by the A N P P (DM) values estimated for these plant life-forms. The % BNPP associated with woody plants was assumed to be the value remaining after subtracting BNPP associated with herbaceous and/or macrophytic plants from the ecosystem BNPP. Estimates of BNPP associated with vascular plant organs were obtained from ecological studies and references as described for the given ecosystem. A l l plant life-form and organ estimates were assembled in a Microsoft Excel spreadsheet containing the ecosystem NPP values from Table 2 . 1 . The Plant Life-Form method was used when the plant NPP information (ANPP and BNPP) was presented separately for individual plant life-forms. Estimates of NPP associated with each plant life-form were obtained from ecological studies and references as described for the given ecosystem. Estimates of BNPP and ANPP associated with vascular plant life-forms were obtained by applying published BNPP:ANPP ratios for the given plant life-form to the NPP associated with the given plant life-form. Estimates of ANPP and BNPP associated with plant organs were obtained in the same manner as for the Belowground-Aboveground method. Some common procedures were followed when using either the Belowground-Aboveground or Plant Life-Form methods. Unless otherwise indicated, only studies that estimated the losses of newly produced plant D M to mortality and/or litterfall of woody plants were used in the estimate. Estimates of the woody plant annual NPP associated with angiosperms and gymnosperms were assumed to be the same for aboveground and belowground plant organs. NPP associated with nonvascular plants was assumed to be completely aboveground. For studies that presented multi-year information for a given site, this information was averaged to represent NPP allocation to organs at the given site. Missing plant life-form and plant organ information was accounted for as noted in the individual ecosystem descriptions. Finally, estimates were presented to one or two decimal places to account for the NPP associated with all plant life-forms and organs. Due to the global nature of this estimate, the reader is advised to consider with caution any meaning in values placed after the decimal point. FOREST. The forest ecosystem is dominated by woody plants that form a closed-canopy. Light penetration to the understory is minimal hence NPP associated with life-forms other than woody plants is small. Forests are located in areas that provide sufficient growth conditions, including mineral nutrition, temperature, and water status, to sustain the growth of woody trees. The Belowground-Aboveground method was employed to estimate plant partitioning of NPP among organs in this ecosystem (Table 2.3). FOREST: Tropical Humid. The combination of nearly constant precipitation (less than two dry months), humid air, and warm climate (mean annual temperature 28°C) creates an environment conducive to continual plant growth, rapid turnover of organic matter, and high rates of net primary productivity in the tropical humid forest (Anonymous 1978a; Brown and Lugo 1982). The ecosystem B N P P A N P P ratio was estimated by Jordan and Escalante (1980) at 1.09. Litterfall measurements were used as a guide to estimate the contribution of vascular epiphytes to the total ANPP. The average rate of vascular epiphyte litterfall (7.0 gnr^y- 1 , mean of 5.5, Nadkarni and Matelson 1992a; 18.0, 0.4, 3.4, 6.0, Tanner, 1980; 11.0, 4.4 Veneklaas 1991) was estimated from studies that measured primarily the litterfall of herbaceous aboveground organs. This value was adjusted for D M loss to retranslocation and herbivory as for woody leaf litter (see below). The corrected value (8.4 g-m"2-y_1) w a s multiplied by 4 to account for woody plant epiphyte ANPP (assuming vascular epiphyte NPP was 25% herbaceous and 75% woody). The final value (33.6 g m ^ - y 1 ) was divided by the ecosystem average rate of net primary productivity (2300 g m " 2 y _ 1 ) estimated by Ajtay et al. (1979) giving the estimated contribution of vascular epiphytes to the ANPP (1.5%). This is a conservative estimate, as Nadkarni and Matelson (1992a) acknowledged problems when sorting epiphyte from woody tree litterfall. Woody epiphytes were assumed to be 100% angiosperm based upon the floristic descriptions by Kress (1989). Estimates of woody and herbaceous epiphyte partitioning of NPP to aboveground and belowground organs were assumed to be the same as those estimated for the terrestrially rooted angiosperm woody and herbaceous plants, respectively, in this ecosystem. o o 00 S, re 9 2 w jo O l 8-8 c o o x x x x X X X X X X X X X X X X X s> s> s> s> n n n n r S X X X X X X X X X ^ >^ O OO O 2 2 2 ; o 5 a m 5 « H T j £ ? d f f l C ^ ^ ^ m w CD CD CD ™ ™ 5° jo oo p 4^ 1 1 1 1 1 p p o © TJ to Ul to to OS l o Ul 4^ . LO b o 1 1 1 1 I I I I 1 1 - J Ul b bo ON Ul to Ul O to OS o to ^ 1 1 1 1 ! y> to p Ul 4* VO O N Ul to to o oo U) o Ul i 4*. 1 1 to 1 4^ I 4*. 1 ON 1 4^ 1 1 I 1 ! 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I I bo Lo Lo 4 ^ ON VO Lo b bo 1 1 LO I 1 1 i t 1 1 i i 1 1 J 1 1 1 1 O o t o VO 1^ oo o t o t o NO U l p 1 1 tO 1 1 I 1 1 1 1 1 1 I 1  t t 1 i i I bo bo t o NO NO b 4 ^ U l 1 1 LO I 1 1 i i 1 1 i i 1 t I t 1 1 1 p o t o _ NO 1^ 00 LO ON U l t o LO 1 1 t o 1 1 1 1 1 1 1 I I 1 1 1 I ! 1 I bo bo io NO bo b bo 1 I U i t o oo p ON U l 1 l 1 1 4 ^ ON LO t o LO LO t o 00 t o OO ON U l U l 1 b NO NO NO U l I I 1 1  1 I 1  ! 1 ! ! Lo Lo NO NO NO NO ON b NO ON LO O 4^  LO Lo 4 . so I I I ! I I CD Cu r—^  cr T 3 81 ANPP associated with nonvascular plants was estimated to be 1.1% by dividing the annual rate of nonvascular plant litterfall production (24.5 g-m^-y 1 , mean of 6.5, 20, Kunkel-Westphal and Kunkel, 1979; 38, Nadkarni and Matelson, 1992a; 53.3, Songwe et al. 1988; 11, 18.4, Veneklaas 1991) by the ecosystem average rate of net primary productivity (2300 gm" 2 y" J) estimated by Ajtay et al. (1979). Estimation of ANPP associated with terrestrially rooted herbaceous plants was based upon the study by Proctor et al. (1983a). In this study, the biomass of terrestrially rooted herbaceous plants was comparable to the biomass of epiphytes; therefore, ANPP associated with terrestrially rooted herbaceous plants was assumed to be 1.5%. Herbaceous plant NPP allocation to aboveground organs was assumed to be the same as that for herbaceous plants in temperate moist grassland. The remaining ANPP (95.9%) was assumed to be associated with terrestrially rooted woody plants, estimated to be 90% angiosperm and 10% gymnosperm based upon the relative absence of gymnosperms from the lowland forests of the Amazon region of South America and Africa (Anonymous 1978&). ANPP associated with reproductive organs of woody plants (angiosperm and gymnosperm) was estimated at 69 g-nr 2-y _ 1 [mean of 123, Haines and Foster 1977; 39, John 1973; 35, Klinge and Rodrigues 1968; 106, 43, Kunkel-Westphal and Kunkel 1979; (79, Foster 1982; 328, Healey and Swift 1977; 60, Smithsonian Environmental Sciences Program 1982; 80, 60, 40, Klinge 1974, 1975; 84, Bernhard 1970; 51, Hladik 1978; 33, Soepadmo 1974, all cited in Leigh, Jr. and Windsor 1982); 33, 82, 26, 56, Lugo and Frangi 1993; 70, Nadkarni and Matelson 1992b; 41, 22, 26, 32, 13, Proctor et al. 1983b; 140, 139, Songwe et al. 1988; 66, 27, Veneklaas 1991)]. ANPP associated with woody plant leaf organs (angiosperm and gymnosperm) was estimated at 798 g-m"2-y1 {mean of leaf litterfall estimates [corrected for retranslocation (8%) and herbivory (9.75%) D M losses as in Songwe et al. (1988)]: 956, 843, Cornforth 1970; 702, Haines and Foster 1977; 892, John 1973; 659, Jordan 1971; 674, Klinge and Rodrigues 1968; 812, 879, Kunkel-Westphal and Kunkel 1979; [735, Foster 1982; 891, Healey and Swift 1977; 772, Smithsonian Environmental Sciences Program 1982; 939, 903, 807, Klinge 1974, 1975; 1048, 915, 855, 753, Bernhard 1970; 783, Hladik 1978; 783, Soepadmo 1972, 1973, 1974, all cited in Leigh and Windsor, 1982], 939, Medina and Zelwer 1972; 573, Nadkarni and Matelson 1992b; 845, Nye 1961; 795, 759, 650, 674, 879 Proctor et al. 1983b; 1086, 1025, Songwe et al. 1988; 591, 659, 525, 662, Tanner 1980; and 555, 340, Veneklaas 1991} and leaf NPP estimates: (950, Bartholomew et al. 1953, cited in Anonymous 1978c; 1200, Kira et al. 1964). ANPP associated with woody plant stem and branch organs was estimated at 1048 g-m-^ y-1 [mean of 1920, Bartholomew et al. 1953, cited in Anonymous 1978b; (700, Rozanov and Rozanova 1964; 900, 750, Muller and Nielsen 1965; 400, Anonymous 1975, all cited in Jordan 1983); 1620, Kira et al. 1964]. In summary, ANPP associated with woody plants (1915 g-m"2-y"') was estimated at 3.6% for reproductive, 41.7% for leaf, and 54.7% for woody organs, the latter of which was subdivided between angiosperm stems (47.6%) and branches (52.4%) and gymnosperm stems (75.7%) and branches (24.3%) based upon the estimates for woody plants from tropical seasonal forest. Estimates of BNPP associated with herbaceous and woody plant organs was based upon the BNPP:ANPP ratio for herbaceous plants in tropical seasonal forest (0.69). BNPP associated with herbaceous plant organs was assumed to be the same as that for herbaceous plants in temperate moist grassland. BNPP associated with woody plant organs was assumed to be the same as that for woody plants in tropical seasonal forest. FOREST: Tropical Seasonal. Tropical seasonal forest is distinguished from tropical humid forest by the lower and more variable levels of precipitation (500-3000 mm, Anonymous 1978a), most of which occurs during the rainy season (Singh and Singh 1993). Hence, the growth period is seasonal and occurs over a period of 4-10 months (Anonymous 1978a). The most obvious changes in the plant life-forms compared to tropical humid forest are the decrease in epiphyte abundance and the increase in terrestrially rooted herbaceous plants. The only estimates of BNPP: ANPP ratios for tropical seasonal forests were obtained using techniques that have been acknowledged to underestimate the BNPP: ANPP ratio (Singh and Singh 1991; Singh and Singh 1993). For this reason, the ecosystem ratio of BNPP:ANPP was assumed to be the same as that for tropical humid forest (1.09). Accounting for the decreased abundance of epiphytes in tropical seasonal forest compared with tropical humid forest, ANPP associated with vascular epiphytes was estimated at 1%. ANPP associated with vascular epiphyte life-forms and organs was assumed to be the same as that for vascular epiphytes in tropical humid forest. ANPP associated with nonvascular plants was estimated to be 0.5% given the less prominent role of mosses in this ecosystem compared to tropical humid forest (Anonymous 1987&). ANPP associated with herbaceous plants was estimated at 9.3% [mean of 11.5, 15.1, 8.9, Chaturvedi and Singh 1987, 11.3, Rana et al. 1988; 6.8 (sal old growth forest), 4.8 (sal seedling coppice forest), Rana et al. 1989; 7.9, 9.2, 9.5, Singh and Singh 1991; 8.4, Singh and Singh 1993]. Herbaceous plant NPP allocated to aboveground organs was assumed to be the same as that for herbaceous plants from temperate moist grassland. The remaining ANPP (89.2%) was assumed to be associated with terrestrially rooted woody plants. Considering the higher gymnosperm content compared to tropical humid forest, 80% and 20% of the woody plant NPP was estimated to be angiosperm and gymnosperm in origin, respectively. ANPP associated with angiosperm woody plants was estimated at 3.6% reproductive (value from tropical humid forest) and the balance at 43.84% for leaf (mean of 40.7, 37.8, Rana et al. 1989; 37.66, 57.1, 42.9, Singh and Singh 1991; 46.9, Singh and Singh 1993), 26.73% for stem (mean of 33.1, 40.6, Rana et al. 1989; 24.68, 17.9, 25.7, Singh and Singh 1991; 18.4, Singh and Singh 1993), and 29.43% for branch organs (mean of 26.2, 21.6, Rana et al. 1989; 37.66, 25.0, 31.4, Singh and Singh 1991; 34.7, Singh and Singh 1993). ANPP associated with gymnosperm woody plants was estimated at 2.3% for reproductive (mean of 2.3, 2.5, 2.2, Chaturvedi and Singh 1987) and the balance at 42.4% for leaf (mean of 60.3, 34.0, 38.4, 37.0, Chaturvedi and Singh 1987), 43.6% for stem (mean of 26.4, 51.6, 46.9, 49.3, Chaturvedi and Singh 1987), and 14.0% for branch organs (mean of 13.3, 14.4, 14.7, 13.7, Chaturvedi and Singh 1987). Estimates of BNPP associated with herbaceous and woody plants were based upon the BNPP:ANPP ratio for herbaceous plants (0.69, mean of 0.61, 0.65, 0.49, Joshi et al. 1990; 1.1, 0.62, Rana et al. 1989). BNPP associated with herbaceous plant organs was assumed to be the same as that for herbaceous plants from temperate moist grassland. BNPP associated with woody plant organs was estimated at 87% for fine roots (mean of 83.0, 92.0, 86.2, Singh and Singh 1991 and 86.7, Singh and Singh, 1993) and 13% for coarse roots (mean of 17.0, 8.0, 13.8, Singh and Singh 1991 and 13.3, Singh and Singh 1993). 22 FOREST: Mangal. The mangal ecosystem is dominated by woody plants (mangroves) living in the tropical saltwater swamp environment. The ecosystem BNPP:ANPP ratio was assumed to be the same as that for saltwater marsh/estuary (3.66). NPP associated with vascular epiphytes was assumed to be negligible as Goulter and Allaway (1979) estimated the non-mangal litterfall contribution to total aboveground litterfall in a mangrove stand to be only 0.24%. NPP associated with herbaceous plants was assumed to be negligible as most mangals have no understory other than mangrove seedlings (Tomlinson 1986). NPP associated with nonvascular plants was assumed to be negligible due to the unfavorable growth conditions of the mud (variable salinity levels) and bark substrates (high levels of tannins) (Hutchings and Saenger 1987; Saenger et al. 1977). Angiosperm woody plants (mangroves) were assumed to be the sole contributor to the A N P P in this ecosystem. ANPP associated with mangroves was estimated at 10.8% for reproductive (mean of 10.0, 11.6, Day et al. 1987) and the balance at 43.0% for leaf [mean of 44.1, 42.0 (litterfall values corrected for 8% D M loss as for tropical humid forest), Day et al. 1987] and 57.0% for woody organs [mean of 55.9, 58.0 (including woody biomass increment and litterfall measurements), Day et al. 1987]. ANPP associated with woody organs was estimated at 47.6% for stem and 52.4% for branch as for angiosperm woody plants from tropical humid forest. BNPP associated with woody plant organs was assumed to be the same as that for angiosperm woody plants from freshwater swamp. FOREST: Temperate Evergreen/Coniferous. Temperate evergreen/coniferous forest was distinguished from temperate deciduous/mixed forest primarily by the nature of leaf growth, the former dominated by evergreen woody plants which retain leaves and the latter dominated by deciduous woody plants which shed leaves in autumn (Kozlowski 1971). Ovington (1983) distinguished two temperate evergreen/coniferous forest eco-types, the broad-leaved (semi-sclerophyllous) forest of warmer climates (mean annual temperature 21.5°C, Burgess 1981) and the needle-leafed coniferous forest of cooler climates (mean annual temperature 6.1 °C, Burgess 1981). Information from studies of both eco-types was combined for this estimate. The ecosystem BNPP:ANPP ratio was estimated at 0.84 (mean of 1.12, Jordan and Escalante 1980; 1.11. 0.30, Keyes andGrier 1981). ANPP associated with vascular epiphytes was estimated at 0.5%, half that of tropical seasonal forest, as epiphytes occur in the warmer areas of temperate evergreen/coniferous forest (Kira 1978). ANPP associated with vascular epiphyte organs was assumed to be the same as that for vascular epiphytes from tropical humid forest. ANPP associated with nonvascular plants was estimated at 2.7% (mean of 0.1, 1.6, 0.8, 8.2, Turner and Long 1975). ANPP associated with herbaceous plants was estimated at 5.5% [mean of 10.4 (chir-pine mixed broadleaf forest), 4.0 (chir-pine forest), 5.3 (mixed-oak/chir-pine forest), 6.4 (rianj-dominated mixed-oak forest), 4.4 (tilonj-dominated mixed-oak forest), Rana et al. 1989; 9.8, 6.1, 7.9, Rawat and Singh 1988; 0 (Pseudotsuga menziesii forest), 0.9 (Pinus ponderosa forest), Whittaker and Niering 1975]. Herbaceous plant NPP allocation to aboveground organs was assumed to be the same as that for herbaceous plants from temperate moist grassland. The remaining ANPP (91.3%) was assumed to be associated with terrestrially rooted woody plants, assuming equal contribution from angiosperms and gymnosperms. ANPP associated with angiosperm woody plants was estimated at 2.4% for reproductive (mean of 2.1, Singh and Yadava 1991; 2.8, Whittaker and Niering 1975) and the balance at 42.0% for leaf (mean of 45.2, 49.0, 17.8, Rana et al. 1989; 44.8, 30.3, 48.1, Rawat and Singh 1988; 41.2, Singh and Yadava 1991; 49.3, 52.6, Whittaker and Niering 1975), 27.2% for stem (mean of 24.1, 23.9, 37.5, Rana et al. 1989; 26.5, 35.7, 21.7, Rawat and Singh 1988; 29.4, Singh and Yadava 1991; 24.6, 21.1, Whittaker and Niering 1975), and 30.8% for branch organs (mean of 30.7, 27.1, 44.7, Rana et al. 1989; 28.7, 34.0, 30.2, Rawat and Singh 1988; 29.4, Singh and Yadava 1992; 26.1, 26.3, Whittaker and Niering 1975). ANPP associated with gymnosperm woody plants was estimated at 4.6% for reproductive (mean of 4.8, 4.3, Whittaker and Niering 1975) and the balance at 34.5% for leaf (mean of 27.4, 23.46, Keyes and Grier 1981; 37.0, 44.4, 19.0, Rana et al. 1989; 23.9, 37.3, 24.0, 44.6, Turner and Long 1975; 45.4, 52.7, Whittaker and Niering 1975), 53.7% for stem (mean of 69.9, 72.26, Keyes and Grier 1981; 43.8, 35.6, 58.5, Rana et al. 1989; 69.9, 56.5, 68.8, 48.9, Turner and Long 1975; 37.1, 29.6, Whittaker and Niering 1975), and 11.8% for branch organs (mean of 2.7, 4.38, Keyes and Grier 1981; 19.2, 20.0, 22.5, Rana et al. 1989; 6.2, 6.2, 7.2, 6.5, Turner and Long 1975; 17.5, 17.6, Whittaker and Niering 1975). Estimates of BNPP associated with herbaceous and woody plants was based upon the BNPP:ANPP ratio for herbaceous plants (0.52, mean of 0.59, 0.36, 0.61, Rawat and Singh 1988). Estimates of BNPP associated with herbaceous plant organs were assumed to be the same as those for herbaceous plants from temperate moist grassland. BNPP associated with woody plants was estimated to be 73.7% for fine roots (mean of 86.4, 61.0) and 26.3% for coarse roots (mean of 13.6, 39.0) based upon the study by Keyes and Grier (1981). FOREST: Temperate Deciduous/Mixed. Temperate deciduous/mixed forest is dominated by a mixture of deciduous (primarily angiosperm) and evergreen (primarily gymnosperm) woody plants (Gower and Richards 1990; Rohrig 1991a). This ecosystem occurs mainly in the Northern Hemisphere and experiences the cool temperate climate (mean annual temperature 9.9°C, Burgess 1981) comparable to that of the needle-leafed temperate coniferous forest (see temperate evergreen/coniferous forest). The ecosystem BNPP:ANPP ratio was assumed to be the same as that for temperate evergreen/coniferous forest (0.84). ANPP associated with nonvascular plants was estimated at 0.4% by dividing the rate of net primary productivity of nonvascular plants (5.1 gm" 2 y _ 1 , Dylis 1971) by the average rate of net primary productivity estimated for temperate deciduous/mixed forest (1300 g-m^-y 1, Ajtay et al. 1979). ANPP associated with herbaceous plants was estimated at 1.3% [mean of 3.8, Borong et al. 1987; 1.1 (Abies concolor ravine forest), 0.1 (Pinus ponderosa-Quercus hypoleucoides forest), 0.3 (Populus tremuloides successional forest), Whittaker and Niering 1975]. Herbaceous plant NPP allocation to aboveground organs was assumed to be the same as that for herbaceous plants from temperate moist grassland. The remaining A N P P (98.3%) was assumed to be associated with woody plants, assuming equal contribution from angiosperms and gymnosperms. ANPP associated with angiosperm woody plants was estimated at 4.4% for reproductive (P. tremuloides forest, Whittaker and Niering 1975), and the balance at 26.6% for leaf (mean of 30.4, 30.2, Ando et al. 1977; 23.0, Pastor and Bockheim 1981; 26.3, 25.0, 25.0, 26.3, 26.94, Ruark and Bockheim 1988), 49.3% for stem (mean of 41.2, 43.4, Ando et al. 1977; 44.7, Pastor and Bockheim 1981; 49.6, 53.0, 53.8, 54.3, 54.44, Ruark and Bockheim 1988), and 24.1% for branch organs (mean of 28.4, 26.4, Ando et al. 1977; 32.3, Pastor and Bockheim 1981; 24.1, 22.0, 21.2, 19.4, 18.62, Ruark and Bockheim 1988). ANPP associated with gymnosperm woody plants was estimated at 4.9% for reproductive (A. concolor forest, Whittaker and Niering 1975) and the balance at 42.44% leaf (mean of 37.73, 47.15), 42.22% for stem (mean of 46.73, 37.72), and 15.34% for branch organs (mean of 15.54, 15.13) (Ando et al. 1977). Estimates of BNPP associated with herbaceous and woody plants was based upon the BNPP:ANPP ratio for herbaceous plants (0.79, mean of 0.94, 0.65, 0.79, Ellenberg et al. 1986, cited in Rohrig, 1991b). Estimates of BNPP associated with herbaceous plant organs were assumed to be the same as those for herbaceous plants from temperate moist grassland. BNPP associated with woody plant organs was assumed to be the same as that for woody plants from temperate evergreen/coniferous forest. FOREST: Boreal Coniferous (closed). The boreal coniferous (closed) forest is located primarily south of the transitional boreal/tundra ecotone in a climate characterized by a short, moist growing season and a long, dry winter (Larsen 1980). The mean annual precipitation is low (400-800 mm), a large proportion of which falls as snow during the winter (Kuusela 1992). The dominant life-form is the woody plant. Plant NPP partitioning estimates were based upon information from subalpine coniferous (closed) forests which are similar to boreal coniferous (closed) forest in terms of climate, growth periods, and vegetation characteristics (Collinson 1977, p 166). The ecosystem BNPP:ANPP ratio was estimated at 2.26 (mean of 1.83, 2.69, Grier et al. 1981). ANPP associated with nonvascular plants was estimated at 4.8%, the mean of the values for nonvascular plants in low arctic/alpine tundra [north of boreal coniferous (closed) forest] and temperate coniferous forest [south of boreal coniferous (closed) forest]. ANPP associated with herbaceous plants was estimated at 1.6% [mean of 5.0, 1.1, Grier et al. 1981; 0 (Abies lasiocarpa subalpine forest), 0.1 (Psueudotsuga menziesii-Abies concolor forest), Whittaker and Niering 1975]. Herbaceous plant NPP allocation to aboveground organs was assumed to be the same as that for herbaceous plants from temperate dry grassland. The remaining A N P P (93.6%) was assumed to be associated with woody plants, estimated to be 20.5% angiosperm and 79.5% gymnosperm based upon biomass estimates from Kuusela (1990, page 30). ANPP associated with angiosperm woody plants was estimated at 2.8% for reproductive (mean of 2.4, 3.2) and the balance at 49.65% for leaf (mean of 48.2, 51.1), 24.3% for stem (mean of 25.3, 23.3), and 26.05% for branch organs (mean of 26.5, 25.6) based upon values from angiosperm shrubs (Whittaker and Niering 1975). ANPP associated with gymnosperm woody plants was estimated at 3.4% for reproductive (mean of 0.9, Bringmark 1977; 4.9, 4.5, Whittaker and Niering 1975) and the balance at 35.2% for leaf (mean of 38.3, Bringmark 1977; 23.8, 30.75, Grier et al. 1981; 44.62, 38.54, Whittaker and Niering 1975), 48.9% for stem (mean of 43.8, Bringmark 1977; 59.2, 57.91, Grier et al. 1981; 39.49, 44.2, Whittaker and Niering 1975), and 15.9% for branch organs (mean of 17.9, Bringmark 1977; 17.0, 11.34, Grier etal. 1981; 15.89, 17.27, Whittaker and Niering 1975). Estimates of BNPP associated with herbaceous and woody plants was based upon the BNPP: ANPP ratio for herbaceous plants in temperate dry grassland (2.54). Estimates of BNPP associated with herbaceous plant organs were assumed to be the same as those for herbaceous plants from temperate dry grassland. BNPP associated with woody plants was estimated at 88.5% for fine roots (mean of 92.5, 82.4, Comeau and Kimmins 1987; 84.9, 94.3, Grier et al 1981) and 11.5% for coarse roots (mean of 7.5, 17.6, Comeau and Kimmins 1989; 15.1, 5.7, Grier etal. 1981). FOREST: Boreal Coniferous (open). The boreal coniferous (open) forest, or taiga, is the transitional ecosystem between the boreal coniferous (closed) forest to the south and the low arctic/alpine tundra to the north (Sirois 1992). Progressing north from the boreal coniferous (closed) forest, the forest canopy becomes discontinuous and there is a greater presence of nonvascular plants (Kershaw 1978). The vegetational and climatic characteristics are transitional between the boreal coniferous (closed) forest and the low arctic/alpine tundra (Sirois 1992). The ecosystem BNPP:ANPP ratio was assumed to be the same as that in the boreal coniferous (closed) forest (2.26). ANPP associated with nonvascular plants was estimated at 6.8% (0stbye et al. 1975). ANPP associated with herbaceous plants was estimated at 4.0% [mean of 7.6, Arthur and Fahey 1992; 3.6 (Betula forest), 0stbye et al. 1975; 0.7 (Pinus pondervsa-Pinus strobiformis forest), Whittaker and Niering 1975]. Herbaceous plant NPP allocation to aboveground organs was assumed to be the same as that for herbaceous plants from temperate dry grassland. The remaining A N P P (89.2%) was assumed to be associated with woody plants, estimated to be 57.8% angiosperm and 42.2% gymnosperm, the averages of the values from low arctic/alpine tundra and boreal coniferous (closed) forest. ANPP associated with angiosperm woody plants was estimated at 2.9% for reproductive [mean of values from low arctic/alpine tundra and boreal coniferous (closed) forest] and the balance at 61.1% for leaf, 22.2% for stem, and 16.7% for branch organs based upon the values for angiosperm shrubs (Whittaker and Niering 1975). ANPP associated with gymnosperm woody plants was estimated to be 4.5% for reproductive (mean of 4.1, Arthur and Fahey 1992; 4.9, Whittaker and Niering 1975) and the balance at 44.5% for leaf [mean of 35.94 (litterfall value corrected for retranslocation as in tropical humid forest), Arthur and Fahey 1992; 53.1, Whittaker and Niering 1975], 40.0% for stem (mean of 50.21, Arthur and Fahey 1992; 29.7, Whittaker and Niering 1975), and 15.5% for branch organs (mean of 13.85, Arthur and Fahey 1992; 17.2, Whittaker and Niering 1975). Estimates of BNPP associated with herbaceous and woody plants was based upon the BNPP:ANPP ratio for herbaceous plants from temperate dry grassland (2.54). Estimates of BNPP associated with herbaceous plant organs were assumed to be the same as those for herbaceous plants from temperate dry grassland. BNPP associated with woody plants was estimated at 99.5% for fine roots and 0.5% for coarse roots (Arthur and Fahey 1992). FOREST: Plantation. The forest plantation is a woody plant-dominated ecosystem established primarily for lumber or fuel and secondarily to remediate degraded environments (Brown et al. 1986). The ecosystem NPP partitioning is dependent upon the location of the plantation (temperate or tropical), the tree species established (angiosperm/gymnosperm, fast vs. slow-growing species), the rotation length, the method of establishment (seedling vs. coppice), the land use prior to plantation (reforestation, afforestation, number of previous rotations), and the degree of management such as weeding, pruning, spacing, and nutrition (Savill and Evans 1986; Evans 1992). Net primary production associated with herbaceous weeds was assumed to be the most important variable affecting NPP allocation among plant life-forms in this ecosystem as Bargali et al. (1992) noted decreased ANPP associated with herbaceous plants with age in an unmanaged Eucalyptus plantation. To account for this, the forest plantation ecosystem was subdivided into two eco-types, <10 yr (pre-establishment) and 11 yr and older (established). Evans (1992) estimated that 75.1% and 24.9% of the total area allocated to forest plantations globally was located in temperate and tropical regions, respectively. Based upon the study by Anonymous (1981), cited in Brown et al. (1986) tropical plantations were subdivided as follows: 44.7% and 19.4% to angiosperm and gymnosperm plantations <10 y, respectively and 28.7% and 7.1% to angiosperm and gymnosperm plantations 11 y and older, respectively. Information from United Nations Economic Commission for Europe and Food and Agriculture Organization of the United Nations (1990) was used to estimate the temperate plantation area associated with angiosperms and gymnosperms. Information was used from temperate countries (20 in total) which classified stocked, exploitable, closed, high forests (not coppice) into angiosperm and gymnosperm by age class. Information was used only from countries that subdivided the areas into the age classes used in this estimate. Countries that lumped natural forests with plantations were not included in the estimate. Temperate plantation area was subdivided as follows: 0.4% and 5.6% to angiosperm and gymnosperm plantations <10 y, respectively and 11.2% and 82.8% to angiosperm and gymnosperm plantations 11 y and older. Overall, forest plantation area was partitioned to angiosperm (11.4%) and gymnosperm (9.0%) plantations <10 y and angiosperm (15.6%) and gymnosperm (64.0%) plantations 11 y and older, respectively. The ecosystem BNPP.ANPP ratio was estimated to be 0.96, the average of the BNPP:ANPP ratios from tropical and temperate forests. FOREST: Plantation-(<10 y). Gymnosperm and angiosperm plantations were assumed to have had similar contributions from herbaceous and nonvascular plants during the first 10 years of establishment and were treated identically in this eco-type. ANPP associated with nonvascular plants was estimated to be 5% as bryophytes and lichens may be important components of the flora prior to canopy closure in young forest plantations (Hill 1979; Kormanik 1979). ANPP associated with herbaceous weed plants was estimated at 12.4%, the median of seven values for Eucalyptus plantations ranging from ages 2 to 8 (Bargali et al. 1992). Herbaceous plant NPP allocation to aboveground organs was assumed to be the same as that for herbaceous plants from temperate moist grassland. The remaining A N P P (82.6%) was assumed to be associated with woody plants, estimated to be 55.9% angiosperm and 44.1% gymnosperm from the information given in the forest plantation ecosystem description (see above). ANPP associated with angiosperm woody plants was estimated at 1.8% for reproductive (mean of 0.8, 1.2, 1.5, 5.3, 2.7, Sukardjo and Yamada 1992; 1.1, 1.3, 0.7, 1.3, Bargali et al. 1992) and the balance at 24.9% for leaf (mean of 26.5, 25.2, 24.6, 23.2, 19.7, 27.6, 27.4), 58.5% for stem (mean of 60.4, 62.5, 64.1, 63.4, 62.0, 49.7, 47.3), and 16.6% for branch organs (mean of 13.1, 12.3, 11.3, 13.4, 18.2, 22.7, 25.3) based upon the study by Bargali et al. (1992). ANPP associated with gymnosperm woody plants was estimated at 1.1% for reproductive (based upon the value from gymnosperm forest plantations 11 y and older) and the balance at 32.6% for leaf, 59.6% for stem, and 7.8% for branch organs as in gymnosperm plantations 11 y and older. Estimates of BNPP associated with herbaceous and woody plants was based upon the BNPP:ANPP ratio for herbaceous plants from the herbaceous reproductive crops ecosystem (0.45). Estimates of BNPP associated with herbaceous plant organs were assumed to be the same as those for herbaceous plants from temperate moist grassland. BNPP associated with woody plants was estimated at 80.4% for fine roots and 19.6% for coarse roots, the averages of the values for woody plants from temperate and tropical forests. FOREST: Plantation^11 y and older). Angiosperm and gymnosperm plantations were treated separately to account for the differences in NPP associated with herbaceous plants. A N P P associated with nonvascular plants in angiosperm and gymnosperm plantations was estimated at 3.2% [mean of 3.2, 3.3 (gymnosperm plantation), Turner and Long 1975]. ANPP associated with herbaceous plants in angiosperm plantations was estimated at 2.3%, half the value for ANPP associated with understory plants in the study by Attiwill (1979). ANPP associated with herbaceous plants in gymnosperm plantations was estimated at 1.4% (mean of 1.3, 1.6, Turner and Long 1975), the lower value accounting for minimal undergrowth after canopy closure in gymnosperm plantations (Hill 1979). Herbaceous plant NPP allocation to aboveground organs was assumed to be the same as that for herbaceous plants from temperate moist grassland. The remaining ANPP in angiosperm plantations (94.5%) was assumed to be associated with angiosperm woody plants. ANPP associated with angiosperm woody plants was estimated at 1.8% for reproductive (based upon the values from angiosperm plantation <10 y) and the balance at 23.44% for leaf, 60.64% for stem, and 15.92% for branch organs (Attiwill 1979). The remaining A N P P in gymnosperm plantations (95.4%) was assumed to be associated with gymnosperm woody plants. ANPP associated with reproductive organs was estimated at 1.1% (Yamakura et al. 1972, cited in Saito 1977) and the balance at 32.6% for leaf (mean of 27.5, 37.2, 37.2, Turner and Long 1975; 28.5, Yamakura et al. 1972, cited in Saito 1977), 59.6% for stem (mean of 65.4, 55.5, 55.7, Turner and Long 1975; 61.9, Yamakura et al. 1972, cited in Saito 1977), and 7.8% for branch organs (mean of 7.1, 7.3, 7.1, Turner and Long 1975; 9.6, Yamakura et al. 1972, cited in Saito 1977). BNPP associated with herbaceous and woody plants and organs was assumed to be the same as that for herbaceous and woody plants from forest plantation <10 y, respectively. TEMPERATE WOODLAND. The temperate woodland ecosystem was considered to be the temperate equivalent of savanna, the woody plant-dominated tropical ecosystem that is transitional between forest and grassland. The Belowground-Aboveground method was employed to estimate plant partitioning of NPP among organs in this ecosystem (Table 2.3). The ecosystem BNPP:ANPP ratio was assumed to be the same as that for seasonal savanna (1.16). ANPP associated with nonvascular plants was estimated at 0.4% as for nonvascular plants from temperate deciduous/mixed forest. ANPP associated with herbaceous plants was estimated at 10.4% [mean of 0.9, {Pinus chihuahuana-Quercus arizonica woodland), 19.9 (Open oak woodland), Whittaker and Niering 1975]. Herbaceous plant NPP allocation to aboveground organs was assumed to be the same as that for herbaceous plants from temperate moist grassland. The remaining A N P P (88.6%) was assumed to be associated with woody plants, assuming an equal contribution from angiosperms and gymnosperms as in the case of temperate forests. ANPP associated with angiosperm woody plants was estimated at 4.4% for reproductive [mean of 3.8, Harcombe et al. 1993; 2.3 (leaf value corrected for retranslocation as in tropical humid forest), Hughes 1971; 7.0 (Open oak woodland), Whittaker and Niering 1975] and the balance at 39.6% for leaf [mean of 37.9 (litter value corrected for retranslocation as in tropical humid forest), Harcombe et al. 1993; 41.23, Whittaker and Niering 1975], 36.6% for stem (mean of 47.6, Harcombe et al. 1993; 25.64, Whittaker and Niering 1975), and 23.8% for branch organs (mean of 14.5, Harcombe et al. 1993; 33.13, Whittaker and Niering 1975). ANPP associated with gymnosperm woody plant organs was assumed to be the same as that for gymnosperm woody plants from temperate deciduous/mixed forest. Estimates of BNPP associated with herbaceous and woody plants was based upon the BNPP:ANPP ratio for herbaceous plants from temperate moist grassland (1.01). Estimates of BNPP associated with herbaceous plants were assumed to be the same as those for herbaceous plants from temperate moist grassland. BNPP associated with woody plants was assumed to be the same as that for woody plants from seasonal savanna. CHAPARRAL, MAQUIS, AND BRUSHLAND. Characterized by a Mediterranean-type climate, with cool, wet winters and hot, dry summers, this ecosystem is dominated by evergreen, sclerophyllous-leaved xerophilous woody plants (Miller and Hajek 1981; Steward and Webber 1981) with or without herbaceous vegetation (Di Castri 1981). The Belowground-Aboveground method was employed to estimate plant partitioning of NPP among organs in this ecosystem (Table 2.3). The ecosystem BNPP:ANPP ratio was estimated at 1.97 (mean of 1.05, 2.89, Kummerow et al. 1981). A N P P associated with nonvascular plants was assumed to be the same as that for nonvascular plants from temperate deciduous/mixed forest (0.4%). ANPP associated with herbaceous plants was estimated at 8.0% (mean of 1.4, 14.7, Kummerow et al. 1981). Herbaceous plant NPP allocation to aboveground organs was assumed to be the same as that for herbaceous plants from temperate dry grassland. The remaining A N P P (91.6%) was assumed to be associated with woody plants, estimated to be 50% angiosperm and 50% gymnosperm based upon ecosystem descriptions by Tomaselli (1981). ANPP associated with angiosperm woody plants was estimated at 8.8% for reproductive (mean of 9.3, 8.4, Kummerow et al. 1981) and the balance at 71.4% for leaf (mean of 74.3, 68.5) and 28.6% for woody organs (mean of 25.7, 31.5) based on the study by Kummerow et al. (1981). ANPP associated with woody organs was estimated at 50.5% for stem and 49.5% for branch as for angiosperm woody plants in seasonal savanna. ANPP associated with gymnosperm woody plants was assumed to be the same as that for gymnosperm woody plants from temperate deciduous/mixed forest. Estimates of BNPP associated with herbaceous and woody plants was based upon the BNPP:ANPP ratio for herbaceous plants from temperate dry grassland (2.54). Estimates of BNPP associated with herbaceous plant organs were assumed to be the same as those for herbaceous plants from temperate dry grassland. BNPP associated with woody plants was estimated at 95.9% for fine roots (mean of 93.8, 98.0) and 4.1% for coarse roots (mean of 6.2, 2.0) based upon the study by Kummerow et al. (1982). SAVANNA. Savannas are open canopy, tropical ecosystems with vegetation characteristics transitional between grassland and woodland. The dominant life-form savanna is the herbaceous plant, the abundance of which is highly sensitive to fire (Singh 1993) and grazing (Pandey and Singh 1992). Ajtay et al. (1979) subdivided savannas into four eco-types based upon physiognomy [see Cole (1963)]. It was not possible to obtain consistent definitions for these eco-types among published studies; therefore, we adapted the savanna classification of Ajtay et al. (1979) to that of Sarmiento and Monasterio (1975) which used climatic and edaphic characteristics to identify three primary savanna eco-types: nonseasonal, seasonal, and hyperseasonal. The Belowground-Aboveground method was employed to estimate plant partitioning of NPP among organs in the three savanna eco-types (Table 2.3). SAVANNA: Nonseasonal. Nonseasonal savannas experience the ever-wet climate of the tropical humid forest; however, the soils are very poor and have low water-retention capacity, leading to excessive draining and the inability to maintain closed-canopy forest (Sarmiento and Monasterio 1975). In this estimate, nonseasonal savannas were substituted for the low tree/shrub savannas identified by Ajtay et al. (1979). The BNPP:ANPP ratio for nonseasonal savanna was assumed to be the same as that from seasonal savanna (1.16). A N P P associated with nonvascular plants was assumed to be the same as that for nonvascular plants from tropical humid forest (1.1%). ANPP associated with herbaceous plants was estimated at 85.4% (mean of 96.4, 90.1, 85.2, 69.7, Menaut and Cesar 1979). Herbaceous plant NPP allocation to aboveground organs was assumed to be the same as that for herbaceous plants from temperate moist grassland. The remaining A N P P (13.5%) was assumed to be associated with woody plants, estimated to be 95% angiosperm and 5% gymnosperm based upon literature descriptions of savanna woody vegetation (Cole 1986). ANPP associated with angiosperm and gymnosperm woody plants was assumed to be the same as that for woody plants from seasonal savanna and temperate deciduous/mixed forest, respectively. Estimates of BNPP associated with herbaceous and woody plants was based upon the BNPP: ANPP ratio for herbaceous plants (0.96, mean of 1.28, 1.04, 0.65, 0.85, Menaut and Cesar 1979). Estimates of BNPP associated with herbaceous plant organs were assumed to be the same as those for herbaceous plants from temperate moist grassland. BNPP associated with woody plant organs was assumed to be the same as that for woody plants from seasonal savanna. SAVANNA: Seasonal. Seasonal savannas experience periods of wet and dry throughout the year. The soils are poor and well-drained (Sarmiento and Monasterio 1979). The seasonal savanna ecosystem was substituted for the dry savanna thorn forest and dry thorny shrub eco-types of Ajtay et al. (1979). The BNPP:ANPP ratio for seasonal savanna was estimated to be 1.16 by Scholes and Walker (1993). A N P P associated with nonvascular plants was assumed to be the same as that for nonvascular plants from temperate deciduous/mixed forest (0.4%). ANPP associated with herbaceous plants was estimated at 36.8% (mean of 35.7, Scholes and Walker 1993; 55.5, 76.0, 53.2, Pandey and Singh 1992; 29.2, 18.1, Grier et al. 1992). Herbaceous plant NPP allocation to aboveground organs was assumed to be the same as that for herbaceous plants from temperate moist grassland. The remaining A N P P (62.8%) was assumed to be associated with woody plants, estimated to be 95% angiosperm and 5% gymnosperm as for woody plants from nonseasonal savanna. A N P P associated with angiosperm woody plants was estimated at 3.6% for reproductive and the balance at 57.4% for leaf and 42.6% for woody organs based upon the study by Scholes and Walker (1993). ANPP associated with woody organs was estimated at 50.5% for stem and 49.5% for branch (Cerocarpus breviflorus shrubland, Whittaker and Niering 1975). A N P P associated with gymnosperm woody plants was assumed to be the same as that for gymnosperm woody plants from temperate deciduous/mixed forest. Estimates of BNPP associated with herbaceous and woody plants was based upon the BNPP:ANPP ratio for herbaceous plants (2.07, Scholes and Walker 1993). Estimates of BNPP associated with herbaceous plant organs were assumed to be the same as those for herbaceous plants from temperate moist grassland. BNPP associated with woody plant organs was estimated at 97.8% for fine roots and 2.2% for coarse roots (Scholes and Walker 1993). SAVANNA: Hyper seasonal. Hyperseasonal savannas experience extended wet and dry periods which expose the vegetation to both extremes of water stress, positive and negative (Sarmiento and Monasterio 1975). The soils are poor, i l l drained, and heavy. The hyperseasonal conditions preclude the growth of most woody plants (Menaut and Cesar 1979); therefore, the hyperseasonal savanna eco-type was substituted for the grass-dominated savanna of Ajtay et al. (1979). The BNPP:ANPP ratio for nonseasonal savannas was estimated to be 1.45 (mean of 1.59, 1.43, 1.32, Menaut and Cesar 1979). A N P P associated with nonvascular plants was assumed to be the same as that for nonvascular plants from seasonal savanna (0.4%). ANPP associated with herbaceous plants was estimated at 95% based upon written descriptions of this eco-type (Menaut and Cesar 1979). Herbaceous plant NPP associated with aboveground organs was assumed to be the same as that for herbaceous plants from temperate dry grassland. The remaining ANPP (4.6%) was assumed to be associated with woody plants, assumed to be 95% angiosperm and 5% gymnosperm as for woody plants from seasonal savanna. ANPP associated with angiosperm and gymnosperm woody plant organs was assumed to be the same as that for woody plants from seasonal savanna. Estimates of BNPP associated with herbaceous and woody plant organs was based upon the BNPP:ANPP ratio for herbaceous plants (0.97, Billore and Mall 1977). BNPP associated with herbaceous and woody plant organs was assumed to be the same as that for herbaceous and woody plants from temperate dry grassland and seasonal savanna, respectively. TEMPERATE GRASSLAND. Temperate grasslands are dominated by herbaceous plants (Coupland 1992). The principle difference between temperate moist and dry grasslands is the annual precipitation which ranges from 750 to 1000 mm in the former and 250 to 500 mm in the latter (Lauenroth 1979). The Plant Life-Form method was employed to estimate plant partitioning of NPP among organs in temperate grasslands (Table 2.4). NPP associated with nonvascular plants in temperate grasslands was estimated at 1% as nonvascular plants are a small component of grassland ecosystems (Coupland 1992). The remaining NPP (99%) was assumed to be associated with herbaceous plants. The BNPP:ANPP ratio for herbaceous plants in temperate moist grassland was estimated at 1.01 [mean of 1.43 (Bridger), 1.63 (Osage), Sims and Singh 1978; 0.35, 0.43, Midorikawa 1972, cited in Midorikawa et al. 1975; 1.22, Kucera et al. 1967]. The BNPP A N P P ratio for herbaceous plants in temperate dry grassland was estimated at 2.54 [mean of 2.25 (Cottonwood), 2.66 (Dickinson), 2.93 (Hays), 2.03 (Hornada-1971), 2.49 (Pantex), 3.37 (Pawnee), Sims and Singh 1978; 2.04, Yano and Kayama 1975]. ANPP associated with herbaceous plants from temperate moist grassland was estimated at 3.6% for reproductive (mean of 6.4, 0.7, Midorikawa 1972, cited in Midorikawa et al. 1975) and the balance at 46.2% for leaf (mean of 46.5, 46.0) and 53.8% for stalk organs (mean of 53.5, 54.0) based upon the study by Midorikawa (1972), cited in Midorikawa et al. (1975). 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C ^ §1 *| o ^ P 3 TJ CD TJ ss. 3 3 P p ' CD 3 Cu 2. vT r r C 0 P P 1—1 cn i_i <*> §s. Cu Cu C <3 CD % o Er 3 U c p 3 OQ 9e grassland was assumed to be 3.6% as in temperate moist grassland, and the balance was estimated at 49.1% for leaf and 50.9% for stalk organs based upon the study by Yano and Kayama(1972). BNPP associated with herbaceous plants in temperate moist grassland was estimated at 40.6% for fine roots (mean of 54.4, 26.8) and 59.4% for coarse roots/rhizomes (mean of 45.6, 73.2) based upon the study by Midorikawa (1972), cited in Midorikawa et al. (1975). BNPP associated with herbaceous plants in temperate dry grassland was estimated at 55.7% for fine roots and 44.3% for coarse roots/rhizomes based upon the study by Yano and Kayama (1975). TUNDRA. Plant growth in the tundra, characterized by an extreme climate of cold (atmospheric temperatures rarely exceeding 10°C) and dry conditions (mean annual precipitation <200 mm), is limited to a short season of <2 months when the permafrost soils are partially thawed (Barry et al. 1981). The dominant life-forms change along a continuum from high latitude/altitude to low latitude/altitude areas. From the former to the latter, nonvascular plants become less dominant and woody plants become more prominent (Webber 1974). Information for estimation of plant NPP partitioning in this ecosystem was obtained from studies that did not completely describe the methods for estimating woody plant NPP. To avoid using information from studies that estimated NPP from annual biomass changes, only studies that indicated the use of correction factors in the results sections were used in this estimate. The Belowground-Aboveground method was employed to estimate plant partitioning of NPP among organs in tundra eco-types (Table 2.3). TUNDRA: Polar Desert. The ecosystem BNPP A N P P ratio was estimated at 0.06 (Bliss 1974). ANPP associated with nonvascular (71.4%) and herbaceous plants (28.6%) was estimated at 71.4% and 28.6%, respectively (Bliss 1974). The BNPP:ANPP ratio of herbaceous plants was estimated at 0.20 (Bliss 1974). ANPP associated with herbaceous plants was assumed to be 1% reproductive to account for the large proportion of sterile herbaceous plants (Alexandrova 1970; Muc 1977), and the balance was assumed to be associated with herbaceous plant organs as for herbaceous plants from high arctic/alpine tundra. BNPP associated with 38 herbaceous plant organs was assumed to be the same as that for herbaceous plants from high arctic/alpine tundra. TUNDRA: High Arctic/Alpine. The ecosystem BNPP:ANPP ratio was estimated at 1.67 (mean of 1.27, 0.86, 1.59, 1.86, Bliss 1974; 2.12, 2.33, Muc 1977). ANPP associated with nonvascular plants was estimated at 50.3% (mean of 45.3, 69.6, 22.7, 62.1, Bliss 1974; 52.0, Bliss et al. 1977). ANPP associated with herbaceous plants was estimated at 28.6% (mean of 45.3, 30.4, 55.3, 5.2, Bliss 1974; 6.6, Bliss et al. 1977). Herbaceous plant NPP allocation to aboveground organs was assumed to be the same as that for herbaceous plants from polar desert tundra. The remaining A N P P (21.1%) was assumed to be associated with woody plants, estimated to be 95% angiosperm and 5% gymnosperm based upon vegetation descriptions in the book by Bliss et al. (1981). ANPP associated with angiosperm and gymnosperm woody plants was estimated at 3% for reproductive (Shaver and Chapin, III 1991) and the balance at 58.9% for leaf (mean of 60.2, 70.2, 61.7, 43.4) and 41.1% for woody organs (mean of 39.8, 29.8, 38.3, 56.6) based upon the Toolik Lake site information for angiosperm woody plants in the study by Shaver (1986). A N P P associated with angiosperm and gymnosperm woody organs was estimated at 57.1% for stem and 42.9% for branch and 72.1% for stem and 27.9% for branch, respectively, based upon the values for woody plants from boreal coniferous (open) forest. Estimates of BNPP associated with herbaceous and woody plants was based upon the BNPP:ANPP ratio for herbaceous plants (2.75, mean of 2.72, 2.70, 2.84, Muc 1977). BNPP associated with herbaceous plants was estimated at 93.8% for fine roots (mean of 95.0, 92.1, 94.2) and 6.2% for coarse roots/rhizomes (mean of 5.0, 7.9, 5.8) based upon the study by Muc (1977). BNPP associated with woody plant organs was assumed to be the same as that for woody plants from boreal coniferous (open) forest. TUNDRA: Low Arctic/Alpine. The ecosystem BNPP A N P P ratio was assumed to be the same as that for high arctic/alpine tundra (1.67). ANPP associated with nonvascular plants was estimated at 35.9% (mean of 50.0, 40.5, 17.7, 35.3, 0stbye et al. 1975). ANPP associated with herbaceous plants was estimated at 39.8% [mean of 7.4 (lichen heath), 46.4 (wet meadow), 71.0 (dry meadow), 34.5 (Salix shrub), 0stbye et al. 1975]. The remaining ANPP (24.3%) was assumed to be associated with woody plants. Estimates of ANPP and BNPP associated with herbaceous and woody plants and organs were assumed to be the same as those for herbaceous and woody plants from high arctic/alpine tundra. DESERT. Deserts are dominated by woody plants capable of living in the extreme climate characterized by low precipitation (desert and semidesert scrub, 100-350 mm; extreme deserts <100 mm) which is highly variable and unpredictable (Noy-Meir 1973). ANPP associated with herbaceous plants in deserts is dependent upon the irregular large bursts of rainfall (Fischer and Turner 1978) and is thus highly variable. The Belowground-Aboveground method was employed to estimate plant partitioning of NPP among organs in this ecosystem (Table 2.3). DESERT: Desert and Semidesert Scrub. Desert and semidesert scrub is dominated by woody plants approaching almost 50% surface cover (West 1983). The ecosystem BNPP:ANPP ratio was estimated at 3.82 [mean of 2.91, 2.88, Caldwell and Camp 1974; 5.55, 0.91, 1.95, 5.26, Rodin 1977, cited in Walter and Box 1983b; 7.90, Balph 1973, cited in Webb et al. 1983; 3.16 (Curlew Valley, Utah), Fischer and Turner 1978]. A N P P associated with nonvascular plants was estimated at 3.0% (Walter and Box 1983c). ANPP associated with herbaceous plants was estimated at 17.6% [mean of 43.2 (Bouteloua curtipendula desert grassland), 9.3 (spinose-suffrutescent desert scrub), 0.4 (Cercidium microphyllum-Franseria deltoidea desert scrub), Whittaker and Niering 1975]. Herbaceous plant NPP allocation to aboveground organs was assumed to be the same as that for herbaceous plants from temperate dry grassland. The remaining A N P P (79.4%) was assumed to be associated with woody plants, estimated to be 100% angiosperm [negligible contribution from gymnosperms based upon floristic descriptions by Shmida (1985)]. ANPP associated with angiosperm woody plants was estimated at 5.5% for reproductive (mean of 5.7, 4.5, 6.2), and the balance at 41.9% for leaf (mean of 53.10, 35.94, 36.66), 17.6% for stem (mean of 11.3, 25.52, 15.89), and 40.5% for branch organs (mean of 35.6, 38.54, 47.45) based upon the study by Whittaker and Niering (1975). Estimates of BNPP associated with herbaceous and woody plants was based upon the BNPP:ANPP ratio for herbaceous plants from temperate dry grassland (2.54). Estimates of BNPP associated with herbaceous plant organs were assumed to be the same as those for herbaceous plants from temperate dry grassland. BNPP associated with woody plants was estimated at 99% for fine and 1% for coarse roots based upon the study by Caldwell and Camp (1974) in which the majority of the BNPP was associated with fine roots. DESERT: Extreme (sandy, hot and dry). Extreme deserts (sandy, hot and dry and sandy, cold and dry) are largely devoid of vegetation (West 1983). The ecosystem BNPP:ANPP ratio was assumed to be the same as that for desert and semidesert scrub (3.82). ANPP associated with nonvascular plants was assumed to be the same as that for nonvascular plants from desert and semidesert scrub (3.0%). ANPP associated with herbaceous plants was assumed to be 5%, a value less than that for herbaceous plants from desert and semidesert scrub, as Whittaker and Niering (1975) demonstrated decreasing ANPP associated with herbaceous plants with increasing xeric status. Herbaceous plant NPP allocated to aboveground organs was assumed to be the same as that for herbaceous plants from desert and semidesert scrub. The remaining A N P P (92.0%) was assumed to be associated with woody plants, estimated to be 100% angiosperm as for desert and semidesert scrub. ANPP associated with angiosperm woody plant organs was assumed to be the same as that for angiosperm woody plants from desert and semidesert scrub. Estimates of BNPP associated with herbaceous and woody plants and organs was assumed to be the same as those for herbaceous and woody plants from desert and semidesert scrub. DESERT: Extreme (sandy, cold and dry). The ecosystem BNPP:ANPP ratio was estimated at 0.11 (total raised beach area, Bliss 1974). ANPP associated with nonvascular and herbaceous plants was estimated at 38.0% and 19.2%, respectively (Bliss 1974). The remaining ANPP (42.8%) was assumed to be associated with woody plants, assumed to be 95% angiosperm and 5% gymnosperm as for woody plants from high arctic/alpine tundra. Estimates of ANPP and BNPP associated with herbaceous and woody plants and organs were assumed to be the same as those for herbaceous and woody plants from high arctic/alpine tundra. LAKE AND STREAM. Freshwater lakes and streams are distinct ecosystems which differ in terms of their hydrology, morphology, circulation of nutrients and organic matter, and trophic status (Payne 1986; Riemer 1984; Taub 1984; Whitton 1975). In spite of these differences, lakes and streams share a common pool of primary producers, aquatic nonvascular plants (macrophytes and non-macrophytes) and aquatic vascular plants (macrophytes). As most of the studies were from lakes, and as the volume of water in rivers is only 1% of that in lakes globally (Wetzel 1975), differences in the NPP allocation patterns of lakes and streams would have little effect on my estimate. The Plant Life-Form method was employed to estimate plant partitioning of NPP among organs in the lake and stream ecosystem (Table 2.4). NPP associated with nonvascular aquatic plants [91.4%, mean of 91.3 (lake), Allanson et al. 1990, p 223; 92.6 (lake), Best 1982; 90.4 (lake), Kajak et al. 1972] was estimated at 39.4% macrophyte and 60.6% non-macrophyte (Howard-Williams and Allanson 1981). The remaining NPP (8.6%) was assumed to be associated with vascular macrophytes.(mean of 8.7, Allanson et al. 1990, p 223; 7.4, Best 1982; 9.6, Kajak et al. 1972). NPP associated with vascular macrophytes was estimated at 45.8% for emergent (mean of 44.8, Best 1982; 18.7, Howard-Williams and Allanson 1981; 74.0. Kajak et al. 1972) and 54.2% for submergent/floating-leaved (mean of 55.2, Best 1982; 81.3, Howard-Williams and Allanson 1981; 26.0, Kajak et al 1972). The BNPP:ANPP ratio for vascular macrophytes was assumed to be the same as that for vascular macrophytes from freshwater marsh (0.82). A N P P associated with emergent macrophyte organs was assumed to be the same as that for emergent macrophytes from freshwater marsh. A N P P associated with submergent/floating-leaved macrophytes was estimated at 2.0% for reproductive (mean of 1.0, Brock et al. 1983; 2.9, Twilley et al. 1985), 50.8% for leaf (mean of 49.4, Brock et al. 1983; 52.2, Twilley et al. 1985), and 47.2% for petiole/stalk organs (mean of 49.5, Brock et al. 1983; 44.9, Twilley et al. 1985). BNPP associated with vascular macrophytes was estimated at 18.6% for fine roots and 81.4% for coarse roots/rhizomes (Twilley et al. 1985). SWAMP, MARSH, AND ESTUARY. Swamps and marshes/estuaries are globally-distributed forested and nonforested wetlands, respectively, characterized by variable levels of flooding, high productivity, rapid decomposition and recycling of nutrients, and usually the absence of deep peat formation (Mitsch and Gosselink 1993). This ecosystem was partitioned into three eco-types, freshwater swamp, freshwater marsh, and saltwater marsh/estuary to account for differences in salinity and dominant plant life-forms. Freshwater swamps and marshes were allocated 0.481 x 10 1 2 m 2 (Matthews and Fung 1987) of the total area allocated to swamps and marshes by Ajtay et al. (1979), 2 x 10 1 2 m 2 . The balance (1.5 x 10 1 2 m 2) was allocated to saltwater marshes plus 1.4 x 10 1 2 m 2 additional to account for estuaries (Table 5.2, Ajtay et al. 1979). Saltwater swamps (mangals) were treated as forests in this estimate. Assuming areal subdivisions to be representative of NPP associated with eco-types, plant NPP associated with the swamp, marsh, and estuary ecosystem was estimated at 9.5% freshwater swamp, 4.6% freshwater marsh, and 85.9% saltwater marsh/estuary. SWAMP, MARSH, AND ESTUARY: Freshwater Swamp. The freshwater swamp ecosystem is dominated by the woody plant life-form. The Belowground-Aboveground method was employed to estimate plant partitioning of NPP among organs in this ecosystem (Table 2.3). The ecosystem BNPP:ANPP ratio was estimated to be 0.75, a higher value than the preliminary BNPP:ANPP ratio estimated by Burns (1984), 0.46 (mean of 0.25, 0.67) which was based upon information from only the top 30 cm of soil. A N P P associated with vascular epiphytes was assumed to be the same as that for temperate evergreen/coniferous forest (0.5%). NPP associated with vascular epiphyte plant organs was assumed to be the same as that for vascular epiphytes from tropical humid forest. ANPP associated with nonvascular plants was assumed to be the same as that for nonvascular plants from temperate evergreen/coniferous forest (2.7%), assumed to be 50% terrestrial and 50% aquatic plants. Nonvascular aquatic plant ANPP was assumed to be associated with macrophytes and non-macrophytes as for nonvascular aquatic macrophytes and non-macrophytes from lake and stream. ANPP associated with terrestrially rooted herbaceous and macrophytic plants was estimated at 17.4% (mean of 16.5, 18.4, Burns 1984), assumed to be 50% herbaceous and 50% aquatic macrophytes. Herbaceous plant NPP allocation to aboveground organs was assumed to be the same as that for herbaceous plants from temperate moist grassland. Macrophytic plant NPP allocation to emergent and submergent/floating-leaved life-forms and aboveground organs were assumed to be the same as that for macrophytes from lake and stream. The remaining ANPP (78.6%) was assumed to be associated with terrestrially-rooted woody plants, assuming equal association with angiosperms and gymnosperms based upon literature descriptions (Mitsch and Gosselink 1993). ANPP associated with angiosperm woody plants was estimated at 2.9% for reproductive (mean of 1.8, 0.6, Burns 1984; 4.4, 1.7, 1.6, 7.1, Megonigal and Day, Jr. 1988) and the balance at 62.0% for leaf (mean of 56.4, 67.7) and 38.0% for woody organs (mean of 43.6, 32.3) based upon the study by Megonigal and Day, Jr. (1988). ANPP associated with angiosperm woody organs was estimated at 46.9% for stem and 53.1% for branch as for angiosperm woody plants from temperate evergreen/coniferous forest. ANPP associated with gymnosperm woody plants was estimated at 2.9% for reproductive as for angiosperm woody plants and the balance at 54.9% for leaf (mean of 58.0, 51.8) and 45.1% for woody organs (mean of 42.0, 48.2) based upon the study by Megonigal and Day, Jr. (1988). ANPP associated with gymnosperm woody organs was estimated at 82.0% for stem (82%) and 18% for branch as for gymnosperm woody plants from temperate evergreen/coniferous forest. Estimates of BNPP associated with herbaceous, vascular macrophytic and woody plants was based upon the BNPP A N P P ratios for herbaceous (1.01, temperate moist grassland) and vascular macrophytic plants (0.82, freshwater marsh). Estimates of BNPP associated with herbaceous plant organs were assumed to be the same as those for herbaceous plants from temperate moist grassland. Estimates of BNPP associated with macrophytic plant organs were assumed to be the same as those for macrophytic plants from lake and stream. BNPP associated with woody plants was estimated at 54.4% for fine roots and 45.6% for coarse roots (Symbala and Day 1988). 44 SWAMP, MARSH, AND ESTUARY: Freshwater Marsh. Freshwater marshes are eutrophic inland wetlands dominated by emergent macrophytes with variable abundance of submergent/floating-leaved macrophytes and nonvascular plants depending upon the water depth and/or cycle stage (Mitsch and Gosselink 1993). There were no studies that estimated NPP associated with the dominant plant life-forms from freshwater marsh; therefore, plant NPP partitioning assumptions were based upon vegetation descriptions by Mitsch and Gosslink (1993). The Plant Life-Form method was employed to estimate plant partitioning of NPP among organs in this ecosystem (Table 2.4). NPP associated with vascular emergent and submergent/floating-leaved macrophytes was estimated at 50% and 25%, respectively. The BNPP:ANPP ratio for vascular macrophytes was estimated at 0.82 [mean of 0.54 (Belleville Pond), 1.11 (Worden Pond, Hogeland and Killingbeck 1985]. A N P P associated with emergent macrophytes was estimated at 2.8% reproductive [mean of 2.2 (Carex gracilis community), 3.4 (Iris pseudoacorus community), Baradziej 1974] and the balance was assumed to be associated with leaf and stalk organs as for herbaceous plants from temperate moist grassland. ANPP associated with submergent/floating-leaved macrophyte organs was assumed to be the same as that for submergent/floating-leaved macrophytes from lake and stream. BNPP associated with vascular macrophyte organs was assumed to be the same as that for vascular macrophytes from lake and stream. NPP associated with terrestrial nonvascular plants was estimated at 5.0% (mean of 4.2, 5.8, Baradziej 1974), and the remaining NPP (20%) was assumed to be associated with nonvascular aquatic macrophytes, associated with macrophyte and non-macrophyte life-forms as for nonvascular aquatic macrophytes from lake and stream. SWAMP, MARSH, AND ESTUARY: Saltwater Marsh/Estuary. Saltwater marshes (open coastal wetlands) and estuaries (partially enclosed coastal wetlands) are dominated by submergent/floating-leaved macrophytes with a less prominent presence of emergent macrophytes and nonvascular plants (Mitsch and Gosselink 1993). The Plant Life-Form method was employed to estimate plant partitioning of NPP among organs in this ecosystem (Table 2.4). NPP associated with aquatic nonvascular plants was estimated at 27.8% (Bally et al. 1985). Aquatic nonvascular plant NPP was estimated at 38.5% for macrophyte (mean of 49.0, Bally et al. 1985; 28.0, Thorn 1984) and 61.5% for non-macrophyte (mean of 51.0, Bally et al. 1985; 72.0, Thorn 1984). NPP associated with emergent macrophytes was estimated at 13.6% (Bally et al. 1985). The remaining NPP (58.6%) was assumed to be associated with submergent/floating-leaved macrophytes. The BNPP:ANPP ratio for vascular macrophytes was estimated at 5.07 (mean of 4.91, da Cunha Lana et al. 1991; 4.6, Smith et al. 1979; 5.70 (ratio for total marsh, high and low), Valiela et al. 1976). ANPP associated with emergent macrophyte organs was assumed to be the same as that for emergent macrophytes from freshwater marsh. ANPP associated with submergent/floating-leaved macrophyte organs was assumed to be the same as that for submergent/floating-leaved macrophytes from lake and stream. BNPP associated with vascular macrophytes was estimated at 18.5% for fine roots and 81.5% for coarse roots/rhizomes (Valiela et al. 1976). ALGAL BED AND REEF. The algal bed and reef ecosystem is composed of nonvascular aquatic plants. The Plant Life-Form method was employed to estimate plant partitioning of NPP among organs in this ecosystem (Table 2.4). NPP associated with nonvascular aquatic plants was assumed to be 100%. Nonvascular aquatic plant NPP was assumed to be associated with macrophyte and non-macrophyte life-forms as for saltwater marsh/estuary. PEATLAND. Peatlands are peat-accumulating, freshwater wetlands located primarily in the cool boreal areas of the Northern Hemisphere. Peatland was subdivided into two eco-types, bog (92.6%) and fen (7.4%, tussock tundra and forb meadows, Matthews and Fung 1987) based upon vegetation and hydrological descriptions by Mitsch and Gosselink (1993) and surface area estimations by Matthews and Fung (1987). Bogs are low nutrient peatlands that receive water and minerals solely from precipitation; whereas, fens receive water and minerals from precipitation and ground water and are more eutrophic than bogs (Crum 1988; Mitsch and Gosselink 1993). 46 PEATLAND: Bog. Bogs are dominated by woody plants and nonvascular plants, the former with stunted growth in the ombotrophic environment (Mitsch and Gosselink 1993). The Belowground-Aboveground method was employed to estimate plant partitioning of NPP among organs in this ecosystem (Table 2.3). The ecosystem BNPP:ANPP ratio was assumed to be the same as that for freshwater swamp (0.75). ANPP associated with nonvascular plants was estimated at 25.8% [mean of 7.9 (Sike Hil l A) , 21.6 (Sike Hi l l B), 16.7 (Bog Hill) , 8.2 (Bog End), Forrest and Smith 1975; 50.1, 50.6, Grigal et al. 1985]. ANPP associated with herbaceous plants was estimated at 17.0% (mean of mean of 30.9, 21.0, 22.5, 22.2, Forrest and Smith 1975; 2.9, 2.2, Grigal et al. 1985). ANPP associated with reproductive organs of herbaceous plants were estimated at 1.8% (mean of 1.7, 2.0, 1.9, 1.5, all from Eriophorum vaginatum, Forrest and Smith 1975) and the balance to leaf and stalk organs as for herbaceous plants from fen. The remaining ANPP (57.2%) was assumed to be associated with woody plants, assumed to be 20.5% angiosperm and 79.5% gymnosperm as for woody plants from boreal coniferous (closed) forest. ANPP associated with woody plant organs was assumed to be the same as that for woody plants from boreal coniferous (closed) forest. Estimates of BNPP associated with herbaceous and woody plants was based upon the BNPP:ANPP ratio for herbaceous plants from fen (1.30). Estimates of BNPP associated with herbaceous plant organs were assumed to be the same as those for herbaceous plants from fen. BNPP associated with woody plants was assumed to be the same as that for woody plants from freshwater swamp. PEATLAND: Fen. Fens are dominated by nonvascular and herbaceous plants (Crum 1988). The Plant Life-Form method was employed to estimate plant partitioning of NPP among organs in this ecosystem (Table 2.4). NPP associated with terrestrial nonvascular and herbaceous plants was estimated at 63.6% and 36.4%, respectively (Billings 1987, cited in Oechel and Billings 1992). The B N P P A N P P ratio associated with herbaceous plants was estimated at 1.30 [mean of 1.0 (Carex acutiformis), 1.59 (C. rostrata), Konings et al. 1992]. ANPP associated with herbaceous plants was estimated at 1.8% for reproductive based upon the value from bogs and the balance to be 68.5% for leaf [mean of 62.5, 71.8, 71.1 (C. diandra)] and 34.5% for stalk organs (mean of 37.5, 28.2, 28.9) based upon the study by Konings et al. (1992). BNPP associated with herbaceous plants was estimated at 74.0% for fine roots (mean of 75.5, 72.5) and 26.0% for coarse roots/rhizomes (mean of 24.5, 27.5) by Konings et al. (1992). CULTIVATED LAND. The Food and Agriculture Organization yield and area data from 1971-1973 (Food and Agriculture Organization of the United Nations Production Yearbook, 1872, 1975, cited in Ajtay et al. 1979) was employed to subdivide cultivated land into five eco-types, assuming surface area partitioning to be representative of NPP partitioning: herbaceous plant crops {84.2% reproductive [cereal, pulse/vegetable (95% assumed to be herbaceous reproductive crop), and oil]; 3.4% shoot [fiber, some pulse/vegetable (5% assumed to be herbaceous shoot crop), and oil]; 5.6% root/tuber} and woody plant crops [reproductive (6.1%); leaf (0.7%), assuming reproductive crops represented the majority (90%) of the woody crop area]. Estimates of plant NPP associated with fore-crops, after-crops, inter-crops, and volunteer crops were not considered in this estimate since the global practices of these agricultural methods were unknown. CULTIVATED LAND: Herbaceous Reproductive Crops. The Plant Life-Form method was employed to estimate plant partitioning of NPP among organs in this ecosystem (Table 2.4). NPP associated with nonvascular plants was estimated at 0.3% (Kukielska 1973, cited in Wojcik 1979). NPP associated with herbaceous weed (non-crop) plants was estimated at 10% based upon the study by Wojcik (1979) in which NPP associated with herbaceous weed plants was shown to vary between 2.7% in a rye field to 21.9% in a potato field. The BNPP: ANPP ratio associated with herbaceous weed plants was estimated at 0.45 (mean of 0.46, 0.44) based upon the study of 2 herbaceous arable plants (meadow and lucerne leys) by Andren et al. (1990). Herbaceous weed plant NPP allocation to aboveground organs was assumed to be the same as that for herbaceous plants from temperate moist grassland. The remaining NPP (89.7%) was assumed to be associated with herbaceous reproductive crop plants. The BNPP:ANPP ratio associated with herbaceous reproductive crop plants was estimated at 0.28 [mean of 0.27 (barley 0), 0.18 (barley 120), Andren et al. 1990; 0.41, Martin and Puckridge 1982; 0.24, Kukielska 1973, cited in Wojcik, 1979]. ANPP associated with herbaceous reproductive crop plants was estimated at 39.0% for reproductive (mean of 44.7, 47.0, Andren et al. 1990; 30.3, 40.0, 38.1, Bolshakov and Rode 1972, cited in Walter and Box 1983a; 33.6, Kukieleska 1973, cited in Wojcik 1979), and the balance was assumed to be associated with leaf and stalk organs as for herbaceous plants from temperate moist grassland. BNPP associated with herbaceous weed and herbaceous reproductive crop plant organs was assumed to be the same as that for herbaceous plants from temperate moist grassland. CULTIVATED LAND: Herbaceous Shoot Crops. The Plant Life-Form method was employed to estimate plant partitioning of NPP among organs in this ecosystem (Table 2.4). NPP associated with nonvascular and herbaceous weed plants and organs was assumed to be the same as that for nonvascular and herbaceous weed plants from the herbaceous reproductive crops ecosystem. The remaining NPP (89.7%) was assumed to be associated with herbaceous shoot crop plants. The BNPP:ANPP ratio for herbaceous shoot crop plants was assumed to be the same as that for herbaceous reproductive crop plants from the herbaceous reproductive crops ecosystem. Estimates of herbaceous shoot crop plant NPP associated with aboveground and belowground organs were assumed to be the same as those for herbaceous weed plants from the herbaceous reproductive crops ecosystem. CULTIVATED LAND: Herbaceous Root/Tuber Crops. The Plant Life-Form method was employed to estimate plant partitioning of NPP among organs in this ecosystem (Table 2.4). NPP associated with nonvascular and herbaceous weed plants and organs was assumed to be the same as that for nonvascular and herbaceous weed plants from the herbaceous reproductive crops ecosystem. The remaining NPP (89.7%) was assumed to be associated with herbaceous root/tuber crop plants. The B N P P A N P P ratio associated with herbaceous root/tuber plants was estimated at 1.92 (Wojcik 1973, cited in Wojcik 1979). Herbaceous root/tuber plant NPP allocation to aboveground;organs was assumed to be the same as that for herbaceous plants from temperate moist grassland. BNPP associated with root/tuber plants was estimated at 13.6% for fine roots and 86.4% for coarse roots/tubers (Wojcik 1973, cited in Wojcik 1979). CULTIVATED LAND: Woody Reproductive Crops. The Belowground-Aboveground method was employed to estimate plant partitioning of NPP among organs in this ecosystem (Table 2.3). The ecosystem BNPP:ANPP ratio was assumed to be the same as that for forest plantations 11 y and older-A (0.96). ANPP associated with nonvascular plants was estimated to be 3.2% as for nonvascular plants from forest plantations 11 y and older-A. ANPP associated with herbaceous weed plants was assumed to be 4% as for herbaceous weed plants from forest plantations 11 y and older-A. Herbaceous weed plant NPP associated with aboveground organs was assumed to be the same as that for herbaceous weed plants from the herbaceous reproductive crops ecosystem. The remaining ANPP (92.8%) was assumed to be associated with woody reproductive crop plants, assumed to be 100% angiosperm based upon descriptions of woody plant crops by Lanner (1981) and Opeke (1982). ANPP associated with reproductive organs of woody reproductive plants was estimated to be 40% based upon the study by Heim et al. (1979) in which the annual D M production of reproductive organs (apples) was estimated to be 57% [mean of 65.6 (Bristol), 48.4 (Montpellier)]. As the Heim et al. (1979) estimate did not account for NPP losses of leaf, stem, and branch organs associated with mortality or herbivory, the estimate was decreased to 40% for reproductive organs. The balance of the ANPP associated with woody reproductive plants (60%) was assumed to be associated with leaf, stem, and branch organs as for angiosperm woody plants from temperate evergreen/coniferous forest. Estimates of BNPP associated with herbaceous weed and woody plants was based upon the BNPP:ANPP ratio for herbaceous weed plants from the herbaceous reproductive crops ecosystem (0.45). Estimates of BNPP associated with herbaceous weed plant organs were assumed to be the same as those for herbaceous weed plants from temperate moist grassland. BNPP associated with woody plant organs was assumed to be the same as that for woody plants from forest plantations. 50 CULTIVATED LAND: Woody Leaf Crops. The Belowground-Aboveground method was employed to estimate plant partitioning of NPP among organs in this ecosystem (Table 2.3). The ecosystem BNPP:ANPP ratio was assumed to be the same as that for the woody reproductive crops ecosystem (0.96). ANPP associated with nonvascular and herbaceous weed plants and organs was assumed to be the same as that for nonvascular and herbaceous weed plants from the herbaceous reproductive crops ecosystem. The remaining ANPP (92.8%) was assumed to be associated with woody leaf crop plants, assumed to be 100% angiosperm based upon woody crop descriptions by Opeke (1982). ANPP associated with reproductive organs of woody leaf crop plants was estimated at 1 % based upon the study by Barua (1987) which indicated low reproductive production in leaf-pruned woody plants. Magambo and Cannell (1981) indicated that pruning also resulted in greater production of leaf organs and lesser production of stem and branch organs in woody leaf crop plants compared to woody plants in temperate ecosystems. By adjusting the values for leaf and woody organs from temperate evergreen/coniferous forest to account for greater leaf production, the balance of the woody leaf crop plant ANPP (99%) was estimated at 55% for leaf and 45% for woody organs. ANPP associated with woody organs was assumed to be the same as that for angiosperm stem (46.8%) and branch organs (53.1%) from temperate evergreen/coniferous forest. BNPP associated with herbaceous weed and woody leaf crop plants and organs was assumed to be the same as that for herbaceous weed and woody reproductive crop plants from the woody reproductive crops ecosystem. HUMAN AREA. Human area includes parks, managed lands (gardens, cemeteries and golf courses), roads, residential areas, industrial areas, water areas (ponds, canals, reservoirs, etc.), and city centers (Gilbert 1989). The dominant plant life-form varies with the different land uses. The Belowground-Aboveground method was employed to estimate plant partitioning of NPP among organs in this ecosystem (Table 2.3). The ecosystem BNPP:ANPP ratio was assumed to be the same as that for temperate forests (0.84). ANPP associated with nonvascular plants was estimated at 5% as nonvascular plants were shown to be highly successful in urban environments (Seaward 1979). ANPP associated with woody plants was estimated to be 30.1% by relating the mean woody plant canopy cover of 26.0% [mean of 41.7 (city), Dorney et al. 1984; 18, 36, 16, 11, 14 (cities), Marotz and Coiner 1973; 24.1 (park), McBride and Jacobs 1976; 24, 22, 36, 37 (cities), Rowntree 1984; 21.6 (city), Sanders 1983; 37.1 (city), Sanders and Stevens 1984] to the ANPP associated with woody plants in an urban community with a 41.7% canopy cover, 46.9% (Dorney et al. 1984). ANPP associated with woody plants was assumed to be 50% angiosperm and 50% gymnosperm as for woody plants from temperate forests. ANPP associated with angiosperm woody plants was estimated at 3.4% for reproductive and the balance at 34.3% for leaf, 38.25% for stem, and 27.45% for branch organs, the averages of the organ values for angiosperm woody plants from temperate evergreen/coniferous and temperate deciduous/mixed forests. ANPP associated with gymnosperm woody plants was estimated at 4.8% for reproductive and the balance at 38.47% for leaf, 47.96% for stem, and 13.57% for branch organs, the averages of the organ values for gymnosperm woody plants from temperate evergreen/coniferous and temperate deciduous/mixed forests. The remaining ANPP (64.9%) was assumed to be associated with herbaceous plants. Herbaceous plant NPP associated with aboveground organs was assumed to be the same as that for herbaceous plants from temperate moist grassland. Estimates of BNPP associated with herbaceous and woody plants were based upon the BNPP:ANPP ratio of herbaceous plants from temperate moist grassland (1.01). Estimates of BNPP associated with herbaceous plants were assumed to be the same as those for herbaceous plants from temperate moist grassland. BNPP associated with woody plant organs was assumed to be the same as that for woody plants from temperate forests. MARINE. The marine ecosystem includes the open ocean, upwelling zones, and continental shelf (Table 5.2, Ajtay et al. 1979) and is dominated by nonvascular aquatic plants. The Plant Life-Form method was employed to estimate plant partitioning of NPP among organs in this ecosystem. NPP associated with nonvascular aquatic plants (non-macrophytic) was assumed to be 100%. 52 Step 3: Estimation of Plant Organ Net Primary Production Associated with Lignin and Holocellulose Averages of the published values for lignin and holocellulose [percentage of all components of the plant D M (organic and inorganic)], obtained from 27 studies (Table 2.5), were applied to the spreadsheet containing the estimated values of the annual NPP associated with plant organs (g DM) to estimate the portion of the annual NPP allocated by plants to lignin and holocellulose synthesis. Step 4: Estimation of Lignin and Holocellulose Net Primary Production Associated with Carbon The atom %C for holocellulose and lignin plant D M (estimated as described below) was multiplied by the estimated plant NPP (DM) allocated to holocellulose and lignin, respectively, in the various plant organs to give the final estimate of the annual NPP (C) allocated by plants to carbohydrate and phenylpropanoid synthesis. The %C in cellulose, an unbranched polymer of repeating glucose units, was taken to be 44.4 (Dragendorff 1884). Hemicellulose is a branched-chain polymer composed of hexoses and pentoses, the proportions of the two varying between plant species (Tarchevsky and Marchenko 1991). Assuming hemicellulose is all pentose or all hexose, the %C would be 45.5 or 44.4, respectively. There is no known hemicellulose at either of these extremes. The difference between these extremes is only 1% C, and since the true value is likely to be in between the two, the value for cellulose (44.4%) was used for both cellulose and hemicellulose. Lignin is a complex polymer consisting of varying proportions of three monolignol building units having the sinapyl, guiacyl, and/or p-hydroxyphenyl substitution patterns, depending on the type of vascular plant species or tissue, being absent from nonvascular plants (Gross 1979). The three units differ in the degree of methoxylation, sinapyl with 2, guiacyl with 1, and p-hydroxyphenyl alcohol with no additional methoxyl groups. Thus, the %C of the lignin o o o < <: <: . Ct> CD CD ft"! ^ ^ g o o C C C g a a a . & c u ,42 VO t O Ul 2 o a < P cn O c > > H f >§ § p p CD ?' ?' B. 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The %C in lignin (64.4%) was estimated as the average of the values from Fagus silvatica (62.6%), Picea excelsa (65.1%), Medicago sativum (65.4%), and Lycopodium obscurum (65.3%) (Freudenberg 1968). 56 RESULTS A N D DISCUSSION Based on the methodologies outlined in the previous section, it was possible to break down the annual global NPP associated with holocellulose and lignin synthesis in plant life-forms and plant organs. These detailed data are presented in the Appendix. Global Production of Lignin and Holocellulose It was estimated that 14% (12 Pg C - y 1 ) of global NPP is associated with the synthesis of lignin while 34% (29 Pg C - y 1 ) is associated with holocellulose (Table 2.6). Thus, 48% (41 Pg C - y 1 ) of the global net assimilation of C is allocated to the production of lignocellulose (lignin plus holocellulose), the largest component of plant cell walls. This is the first estimate of the annual net flow of C to plant biochemical compounds on a global scale and may serve as a basis for modeling changes in patterns of C allocation as ecosystems respond to climate change. Ecosystem Contribution to Global Net flow of Carbon to Lignocellulose Lignocellulose production comprises 61% (38 Pg C y 1 ) of NPP in terrestrial ecosystems, with holocellulose constituting the majority (Table 2.6). Lignin production does not occur and holocellulose production is comparatively smaller in the marine ecosystem (2.6 Pg C - y 1 , 11% of NPP) (Table 2.6). The dominant ecosystems that were estimated to be associated with 90% of the global net flow of C to lignocellulose are forest (15.6 Pg C-y _ 1 ) , savanna (10.7 Pg C-y 1 ) , cultivated land (3.7 Pg C - y 1 ) , temperate grassland (2.5 Pg C-y - 1 ) , marine (2.6 Pg C - y 1 ) , and swamp, marsh and estuary (2 Pg C-y 1 ) . In the following sections, each ecosystem is treated separately to elucidate the role of lignocellulose in the C cycling processes inherent to the respective ecosystem. Only the dominant plant life-forms and organs that are responsible for the majority of the lignocellulose production are highlighted. Discussion of the relative residence times of the pools of C associated with lignocellulose in plant matter is based upon the lignocellulose content of the Table 2.6 Annual net flow of photosynthetically fixed carbon (C) to global lignin and holocellulose synthesis. [Values in parentheses represent % of the respective net primary production (NPP) category; 1 Pg=101 5 g. Terrestrial, marine, and global total NPP (C) values used to obtain the % values were estimated assuming the conversion factor from dry matter to C is 0.45 (Ajtay et al. 1979)] TERRESTRIAL NPP M A R I N E NPP G L O B A L NPP P L A N T P O L Y M E R (Pg C-y l ) (PgC-y-l) (PgC-y-1) Lignin 12(19) — 12(14) Holocellulose 26 (42) 11(5) 29 (34) Total (Lignocellulose) 38(61) 11(5) 41 (48) plant organs listed in Table 2.5 and the relative rates of plant organ decomposition. Plant organs were classified into one of three categories of lignocellulose content, high (>76% DM), medium (26-75% DM), and low (<25% DM) (Table 2.7). Plant organs were further grouped into relative C turnover class based upon the general finding that the rate of detrital C turnover in terrestrial and marine ecosystems is inversely proportional to the lignocellulose content of plant matter (Herman et al. 1977; Kristensen 1994; Schimel et al. 1994; Stohlgren 1988). Finally, plant organs were grouped according to C-sink potential assuming that the C sink potential is inversely proportional to the rate of C turnover. Although it has been documented that the relative rates of detrital C turnover for the polymers compris ing l ignocel lu lose are hemicellulose>cellulose>lignin (Herman et al. 1977; MacMil lan 1988), holocellulose decomposition has been shown to be tightly linked to that of lignin in the later stages of plant matter decomposition (Berg et al. 1984; Gong et al. 1981; Melillo et al. 1989). Lignocellulose production in the forest ecosystem. Lignocellulose synthesis was highest in the forest ecosystem which was estimated to be associated with 44% (5.1 Pg C-y 1 ) of the global net flow of C to lignin (Fig. 2.2A) and 35% (10.2 Pg C - y 4 ) to holocellulose (Fig. 2.3A). This is not surprising, since in addition to their high total NPP allocation (25.9%, Table 2.1), forests were estimated to be associated with 63% and 78% of the annual NPP (DM) of woody angiosperm and gymnosperm plants, respectively (Fig. 2.4A and B), the life-forms containing the organs with the highest lignocellulose content (Table 2.7). Since the net flow of C to lignocellulose in the forest ecosystem (15.3 Pg C - y 1 ) encompasses 18% of global NPP, knowledge of the ultimate fate of this C in different eco-types is crucial to understanding the patterns of current C sinks and sources in terrestrial ecosystems. Aboveground, the majority of lignocellulose production was associated with woody angiosperm and gymnosperm stem and branch organs (5.4 Pg C - y 1 ) and leaf organs (1.9 Pg C-y 1 ) . Belowground, woody angiosperm and gymnosperm coarse root organs were estimated to be associated with 1.6 Pg C - y 1 and fine root organs with 5.6 Pg C - y 1 . Since roughly 60% of this lignocellulose production was associated with tropical forests (humid and seasonal) and the remainder mainly with forests of 59 Table 2.7 Lignocellulose content, rate of carbon (C) turnover, and C sink potential of plant organs. (A, angiosperm; AQ, aquatic; Above, aboveground; E, emergent; G, gymnosperm; M , macrophyte; non, nonvascular; N M , non-macrophyte; S/FL, submergent/floating-leaved; TE, terrestrial) P L A N T O R G A N LIGNOCELLULOSE CLASS (% Dry Matter) R A T E OF C TURNOVER C-SINK POTENTIAL Above: non-AQ (M, NM) Tuber: herbaceous Low (<25%) High Low Leaf: A l l vascular life-forms Stalk: herbaceous, macrophyte (E) Stalk/Petiole: macrophyte (S/FL) Reproductive: angiosperm Fine root: all vascular life-forms Coarse root/rhizome: herbaceous; macrophyte (E, S/FL) Above: non-TE Medium (26-75%) Medium Medium Stem: woody (A, G) Branch: woody (A, G) Coarse Root: woody (A, G) Reproductive: woody (G) High (>76%) Low High Forest 5.1 P g C - y " 1 (44%) Cultivated Land l . O P g C x 1 (8%) Swamp, marsh, estuary 0.6 Pg C - y 1 (5%) Savanna 3.1 Pg C-y" 1 (27%) Temperate Grassland 0.7 Pg C - y 1 (6%) • LF-Woody (A) Q L F - W o o d y ( G ) U ST/BR-Woody (A) • ST/BR-Woody (G) B F R - W o o d y ( A ) E_ FR-Woody(G) __ C R - W o o d y ( A ) • C R - W o o d y ( G ) __ LF-Herbaceous • • 13 0 ED SK-Herbaceous RE-Herbaceous FR-Herbaceous CR/R/T-Herbaceous LF-Macrophyte SK/P-Macrophyte FR-Macrophyte CR/R-Macrophyte O T H E R ure 2.2 Plant organ contribution to lignin production in forest, savanna, cultivated land, temperate grassland, and swamp, marsh, and estuary ecosystems. Numbers in parentheses represent percentage of global net flow of carbon (C) to lignin. (A, angiosperm; B R , branch; C R , coarse root; FR, fine root; G , gymnosperm; L F , leaf; P, petiole; Pg, petagram=10 1 5g; R, rhizome; R E , reproductive; S K , stalk; ST, stem; T, tuber) 61 Cultivated Land 2.8 P g C - y - 1 (10%) Swamp, marsh, and estuary 1.4 P g C - y 1 (5%) Temperate Grassland 1.8 Pg C - y 1 (6%) LF-Woody (A) LF-Woody (G) ST/BR-Woody (A) ST/BR-Woody (G) FR-Woody (A) FR-Woody (G) CR-Woody (A) CR-Woody (G) LF-Herbaceous • • SK-Herbaceous RE-Herbaceous FR-Herbaceous CR/R/T-Herbaceous LF-Macrophyte SK/P-Macrophyte FR-Macrophyte CR/R-Macrophyte A B O V E - N o n (AQ) O T H E R Figure 2.3 Plant organ contribution to holocellulose production in forest, savanna, cultivated land, temperate grassland, and swamp, marsh, and estuary ecosystems. Numbers in parentheses represent percentage of global net flow of carbon (C) to holocellulose. (A, angiosperm; A Q , aquatic; A B O V E , aboveground; B R , branch; C R , coarse root; FR, fine root; G , gymnosperm; L F , leaf; Non, nonvascular; P, petiole; Pg, petagram=10 1 5g; R, rhizome; R E , reproductive; S K , stalk; ST, stem; T, tuber) 62 Woody Angiosperm 51.4 Pg D M - y - 1 Herbaceous 54.7 Pg D M - y 1 F TW C M B S T G D m SME 19 PL • C L • M • OTHER • n • m B Woody Gymnosperm 18.3 Pg D M - y - 1 Nonvascular ( A Q , N M ) 54.1 Pg D M - y 1 Figure 2.4 Ecosystem contribution to annual net primary production of angiosperm woody, gymnosperm woody, herbaceous, and nonvascular aquatic (non-macrophyte) plant dry matter (DM). ( A Q , aquatic; C M B , chaparral, maquis, and brushland; C L , cultivated land; D , desert; F, forest; M , marine; N M , non-macrophyte; Pg, petagram=10 1 5g; P L , peatland; S, savanna; S M E , swamp, marsh, and estuary; T G , temperate grassland; T W , temperate woodland) the Northern Hemisphere (boreal and temperate), lignocellulose production in the plant organs mentioned above is subdivided into tropical (60%) and Northern Hemisphere (40%) forests. Tropical forests are thus associated with an annual net flow of roughly 4.2 Pg C and 4.5 Pg C, respectively, to high C-sink potential (stem, branch, and coarse root) and medium C sink-potential (leaf and fine root) plant organs. Assuming ecosystems to be in steady state, such that annual NPP is equivalent to mortality plus litterfall and there is no net growth of plant D M , the annual net flow of C to lignocellulose in new plant growth should be equivalent to the annual net flow of C to the detrital pool. This is supported by recent research which documented a positive correlation between primary production and C flux to the detrital pool for a number of terrestrial and aquatic ecosystems (Cebrian and Duarte 1995). Thus, the magnitude of lignocellulose C entering the high and medium C-sink potential plant organ detrital pools as stated above suggests a role for tropical forests as an annual net C sink. Decomposition rates for plant organs in tropical forests are relatively high, with 70% of leaf litter (Pant and Tiwari 1992; Swift et al. 1979, p. 317) and 25% of stem and branch litter (Swift et al. 1979, p. 317) decomposing within one year. Since the relative lignin contents of leaf and fine root organs are similar (Table 2.5) and the rate of lignin decomposition is the main determinant of the rate of plant litter decomposition (see above), fine root decomposition was assumed to be the same as that for leaf organs. Employing a calculation in which the percentage of undecomposed plant organ litter remaining after one year of decomposition is multiplied by the annual C flow to lignocellulose in the given plant organ, a rough estimate of the annual C sink in lignocellulose is obtained. This is only a preliminary estimate since the relative rates of decomposition for different plant organ litter constituents are not equivalent. However, since the lignocellulose polymers are the limiting factors to plant organ decomposition (see above), the following estimates may be representative of the current patterns of lignocellulose deposition and C-sink potential in global ecosystems. Thus, lignocellulose production is an estimated annual sink of 0.3 Pg C y 1 and 1 Pg C - y 1 , respectively, in the medium C-sink potential leaf and fine root organs. The high C-sink potential woody stem and branch organs are an estimated sink of at least 3.4 Pg C - y 1 . Overall, lignocellulose production in undisturbed tropical forests may represent an annual sink of 4.7 Pg C - y 1 . This is twice the estimate of Lugo and Brown (1992) which indicated that tropical forests in 1980 were a potential sink of 1.5-3.1 Pg C - y 1 . However, since 1980, the rate of tropical deforestation has increased by 90%, rendering tropical forests a net source of 1.1-3.6 Pg C - y 1 (Ciais et al. 1995; Houghton 1991). Northern Hemisphere forests are associated with the the majority of the remaining 40% of forest lignocellulose production. Thus, 2.8 Pg C - y 1 and 3 Pg C - y 1 were estimated to be allocated to high C-sink potential (stem, branch, and coarse root) and medium C sink-potential (leaf and fine root) plant organs, respectively. Rates of decomposition in temperate and boreal forests are much slower than in tropical forests (Ciais et al. 1995; Eijsackers and Zehnder 1990; Shaver et al. 1992; Vogt et al. 1986). This is related to the lower mean annual temperature and precipitation of temperate and boreal latitudes. Annual decomposition rates for leaf, fine root, and stem/branch/coarse root organ litters have been estimated at 25%, 20%, and 5%, respectively (Edmonds 1984; MaClaugherty et al. 1982; Stohlgren 1988; Swift et al. 1979, p. 309). Thus, lignocellulose production in medium C-sink potential organs is an estimated sink of 2.3 Pg C - y 1 (0.5 from leaf, 1.8 from fine root organs). Lignocellulose production in high C-sink potential stem/branch/coarse root organs is an estimated sink of 2.7 Pg C - y 1 . The total annual C sink potential in undisturbed Northern Hemisphere forests is thus 5 Pg C - y 1 . This is consistent with recent reports indicating that Northern forests may sequester between 2 to 3.5 Pg C - y 1 (Ciais et al. 1995; Tans et al. 1990). The difference between the two estimates may be attributed to C released to the atmosphere as a result of boreal forest fires (Kasischke et al. 1995). Lignocellulose production in the savanna ecosystem. The savanna ecosystem was second to forests in terms of the magnitude of global lignocellulose production and was estimated to be associated with 27% (3.1 Pg C - y 1 ) of the global net flow of C to lignin (Fig. 2.2B) and 26% (7.4 Pg C - y 1 ) to holocellulose (Fig. 2.3B). This is consistent with their role in the global NPP budget. Savannas accounted for a large amount of global NPP (20.9%, Table 2.1) and were a dominant contributor to the terrestrial NPP of the high lignocellulose-containing angiosperm woody plants (22%, Fig. 2.3 A ; Table 2.7). In addition, savannas were the largest contributor to the annual NPP (DM) of medium lignocellulose-containing herbaceous plants (49%, Figure 2.2C; Table 2.7). Aboveground, high C-sink potential plant organs (woody angiosperm stem and branch) were estimated to be associated with 0.9 Pg C-y" 1. Medium C-sink potential organs (angiosperm leaf and herbaceous leaf and stalk) were associated with 3.8 Pg C - y 1 . Belowground, medium C-sink potential organs (woody angiosperm fine root and herbaceous fine and coarse root/rhizome) were estimated to be associated with 5.6 Pg C-y"1. Since decomposition in nonseasonal savannas is not precipitation-limited and has been shown to be 20 times more rapid than in seasonal and hyperseasonal savannas (Sholes and Walker 1993 p. 183), discussion of lignocellulose production in the savanna ecosystem is subdivided into nonseasonal and seasonal/hyperseasonal. Nonseasonal savannas were estimated to be associated with 16% of the annual net production of lignocellulose and seasonal/hyperseasonal with the remainder. Rates of decomposition for nonseasonal savannas were assumed to be similar to rates in tropical forests. Thus, lignocellulose production in nonseasonal savannas was estimated to be a C sink of 0.1 Pg C-y" 1 in woody stem/branch, 0.2 Pg C-y - 1 in leaf/stalk, and 0.3 Pg C-y"1 in woody angiosperm fine root and herbaceous fine root/coarse root/rhizome organs. Annual rates of plant organ litter decomposition in seasonal and hyperseasonal savannas were estimated to be 10% in woody angiosperm stem/branch organs (assuming a lower rate than nonseasonal savanna due to less precipitation), 100% in leaf/stalk organs (Sholes and Walker 1993, p. 182), and 56% in woody angiosperm fine root and herbaceous fine root/coarse root/rhizome organs (mean of 52, Kaiser 1983; 60, Pandeya et al. 1986, cited in Singh and Gupta 1993). Thus, lignocellulose production in seasonal/hyperseasonal savanns was estimated to be a C sink of 0.7 Pg C - y 1 for woody angiosperm stem/branch, 0 Pg C-y"1 in leaf/stalk, and 2.1 Pg C - y 1 in woody angiosperm fine root and herbaceous fine root/coarse root/rhizome organs. The total C-sink potential associated with lignocellulose production in undisturbed savanna ecosystems is 3.4 Pg C - y 1 . A significant anthropogenic disturbance of savannas is that of biomass burning which has been estimated to release 1.3 Pg C - y 1 into the atmosphere (Ciais et al. 1995). Assuming that savanna fires eliminate lignocellulose C-sinks predominantly from aboveground plant organs, this suggests that the annual lignocellulose production associated with fine roots (2.4 Pg C - y 1 ) may be the only net annual C-sink associated with plant organs in savannas. That the dominant C-sink organs are fine roots is consistent with current research which implicates herbaceous fine roots as potential C sinks in savanna ecosystems (Fisher et al. 1994). Lignocellulose production in the cultivated land ecosystem. The cultivated land ecosystem was estimated to be associated with 8% (1.0 Pg C - y 1 ) of the global net flow of C to lignin (Fig. 2.2C) and 10% (2.8 Pg C - y 1 ) to holocellulose (Fig. 2.3C). The majority of this C flow (90%) was associated with the medium C-sink potential organs of herbaceous plants in the herbaceous reproductive crops eco-type. As noted in Fig. 2.4C, cultivated lands are associated with 26% of the annual NPP of herbaceous plants. Aboveground, herbaceous plant leaf, stalk, and reproductive organs were estimated to be associated with 2.7 Pg C - y 1 in lignocellulose. Reproductive organs represent roughly 30% of the aboveground net flow of C to lignocellulose. Since the reproductive organs are harvested and the leaf and stalk organs are removed following cultivation, lignocellulose production in aboveground organs does not represent a potential C-sink. Belowground, woody angiosperm fine root and herbaceous fine root and coarse root/rhizome/tuber organs were estimated to be associated with 0.8 Pg C-y - 1 in lignocellulose. Assuming the rate of root litter decomposition to be between that of tropical grassland (hyperseasonal savanna) (56%) and temperate grassland (20%), lignocellulose production in belowground organs may represent a small annual sink of between 0.4 and 0.6 Pg C - y 1 . This is consistent with the opinion of Cole et al. (1993) that current agricultural lands are in steady state with the global C budget. Lignocellulose production in the temperate grassland ecosystem. The temperate grassland ecosystem was estimated to be associated with 6% (0.7 Pg C-y 1 ) of the global net flow of C lignin (Fig. 2.2D) and 6% (1.8 Pg C - y 1 ) to holocellulose (Fig. 2.3D). A l l of this C flow was associated with the medium C-sink potential organs of herbaceous plants since temperate grasslands were estimated to be associated with 5% of global NPP (Table 2.1) and 18% of the annual NPP of herbaceous plants (Fig. 2.4C). Aboveground, medium C-sink potential organs (herbaceous leaf and stalk) were estimated to be associated with 1.0 Pg C-y- 1 in lignocellulose. As decomposition of leaf and stalk organ litters in temperate grasslands has been estimated at 25% (Ando 1975), leaf and stalk organs may represent a C sink of 0.8 Pg C - y 1 . Belowground, medium C-sink potential organs (herbaceous fine root and coarse root/rhizome) were estimated to be associated with 1.4 Pg C - y 1 in lignocellulose. Decomposition of roots in temperate grassland was estimated to be roughly 35% less than that in seasonal savannas (Kaiser 1983); therefore, the rate of root litter decomposition is assumed to be 20%. Thus, belowground organs may act as a sink of 1.1 Pg C - y 1 . Combining aboveground and belowground organs, temperate grasslands are estimated to be a potential sink of 1.9 Pg C - y 1 . That temperate grasslands may be a C sink is supported by the Ojima et al. (1993) grassland estimate of 0.6 Pg C - y 1 . Lignocellulose production in the swamp, marsh, and estuary ecosystem. The swamp, marsh, and estuary ecosystem was estimated to be associated with 5% (0.6 Pg C - y 1 ) of the global net flow of C lignin (Fig. 2.2E) and 5% (1.4 Pg C - y 1 ) to holocellulose (Fig. 2.3E). Although the contribution of the swamp, marsh, and estuary ecosystem to the total global NPP (5%) is the smallest of those considered in the previous sections, this is main ecosystem contributing to the annual production of vascular macrophytes. Aboveground, macrophytic plant leaf and stalk organs were estimated to be associated with 0.4 Pg C - y 1 lignocellulose of which greater than 60% occurred in the saltwater marsh/estuary eco-type. Nonvascular aquatic (macrophyte and non-macrophyte) plants were associated with 0.1 Pg C - y 1 , primarily in the saltwater marsh/estuary eco-type. A short-term study by Kristensen (1994) estimated between 29-33% and 40-44% decomposition of vascular macrophyte and nonvascular macrophyte litters, respectively, within 70 days. Thus, aboveground organs do not appear to represent a significant potential C sink in this ecosystem. Belowground, macrophytic plant fine root and coarse root/rhizome organs were estimated to be associated with 1.3 Pg C - y 1 , primarily in the saltwater marsh/estuary eco-type. Based upon the estimate of root decomposition in a saltwater marsh (20%, Hackney and de la Cruz 1980), belowground lignocellulose production was estimated to be a potential sink of 1 Pg C - y 1 . Thus, lignocellulose production in the undisturbed swamp, marsh, and estuary ecosystem may be a net annual sink of 1 Pg C-y- 1. This may change in the future as pollutant run-off from terrestrial ecosystem increases. Lignocellulose production in the marine ecosystem. The marine ecosystem was estimated to be associated with 9% (2.6 Pg C-y 1 ) of the net flow of C to lignocellulose. This C flow was attributed to nonvascular aquatic (non-macrophyte) plants since the marine ecosystem is the main contributor to the global NPP of this life-form (Fig. 2.4D). This amount is small compared to the total annual NPP associated with the marine ecosystem (51.3 Pg DM.y-1, 27.3%) since lignin is not produced by nonvascular plants. The annual rate of nonvascular aquatic plant litter decomposition was estimated to be nearly complete since Taylor et al. (1991) estimated 90% recycling of phytoplankton NPP. Thus, annual lignocellulose production in the marine ecosystem is estimated to be a small potential sink of 0.3 Pg C - y 1 . Lignin Production and the Global Nitrogen Cycle At the cellular level, lignin synthesis interacts with N metabolism at the prephenate-arogenate junction, the metabolic link between the shikimate and phenylpropanoid pathways (Fig. 2.5). Using 15N-phenylalanine, it was recently demonstrated in a number of plant storage organs that when phenylpropanoid metabolism is stimulated to synthesize lignin, the N which is cleaved from phenylalanine is recycled via glutamate through the series of transaminations leading back to the prephenate-arogenate junction (Razal et al. in press). In the presence of the glutamine synthetase inhibitor methionine sulfoximine, however, the ammonium was released and accumulated as such. At the global level, we estimated the annual reassimilation of N through the prephenate-arogenate junction to be l x l O 1 4 mol. The annual net flow of C to lignin synthesis (12 Pg, Table 2.6) was first converted to moles. Since the lignin polymer is composed of 9-carbon building units derived from the phenylpropanoid pathway, for each 9 moles of C incorporated into lignin, 1 mole of N is reassimilated at the prephenate-arogenate junction. Thus, our final estimate was obtained by assuming that one-ninth of the global net flow of C to lignin synthesis was associated with N reassimilation. Using a simpler approach to estimate the annual net flow of C to lignin, Raven et al. (1992) obtained the same estimate of N reassimilation, l x l O 1 4 mol-y-1- In their estimate, global 69 Erythrose-4-phosphate + Phosphoenolpyruvate 1 ^ Shikimate Pathway ^ I ^ Phenylpropanoid Pathway I Lignin Figure 2 . 5 Diagrammatic representation of nitrogen ( N ) reassimilation during lignin biosynthesis at the prephenate-arogenate junction. ( 1 , prephenate aminotransferase; 2 , arogenate dehydratase; 3 , phenylalanine ammonia lyase; 4, glutamine synthetase; 5 , glutamate:oxoglutarate aminotransferase). [Modified from Razal et al (in press)] NPP was assumed to be 95 Pg C - y 1 as opposed to 84.6 Pg C - y 1 (our estimate), the difference due to a higher value of oceanic NPP. As in our estimate, Raven et al. (1992) accounted for differences in phenylpropanoid (lignin) content (% total C) of marine vascular macrophytes (5%), woody plants (20%), and herbaceous plants (<20%). Aside from acknowledging the use of a published estimate of marine vascular macrophyte production, no methods were cited in the estimation of plant life-form contribution to terrestrial NPP. Our estimate overcomes this limitation by providing detailed information on the methodologies used to partition NPP among plant life-forms and plant organs. Our estimate is an improvement of the Raven estimate as it provides additional information on the global breakdown of N reassimilation associated with plant life-forms and plant organs in 37 ecosystems. This information is readily available through the simple conversion of the C allocated to lignin synthesis per plant organ (Fig. 2.2; Appendix) to N reassimilation as described above. The trends of global N reassimilation should be identical to those presented in Fig. 2.2 for lignin production, with forest the dominant ecosystem, followed by savanna, cultivated land, temperate grassland, and swamp, marsh, and estuary. Since global warming is expected to lead to an increase in the C:N ratio of plant D M (Rastetter et al. 1992), which is predicted to increase ecosystem C storage and decrease N availability, and plant growth is already N-limited in terrestrial (Tamm 1991) and marine ecosystems (Cullen et al. 1992), this information will be critical to modeling future changes to plant growth. Allocation of Global Net Primary Production Plant life-forms. Based upon the 160 references and the methodologies explained in the methods section of this chapter, global allocation of NPP (DM) among plant life-forms was estimated at 29% for herbaceous, 29% for nonvascular aquatic (non-macrophyte), 27% for angiosperm woody, 10% for gymnosperm woody, 3% for macrophyte (78% of which is submergent/floating-leaved), 1% for nonvascular terrestrial, and 1% for nonvascular aquatic (macrophyte) (Fig. 2.6). Such a comprehensive estimate based upon published data and Figure 2.6 Anual net primary production (NPP) associated with plant life-forms and plant organs. (A, angiosperm; A B O V E , aboveground; A Q , aquatic; B R , branch; C R / R / T , coarse root/rhizome/tuber; D M , dry matter; F R , fine root; G , gymnosperm; L F , leaf (including petiol); M , macrophyte; N M , non-macrophyte; R E , reproductive; S K / P (petiole for macrophyte-submergent/floating-leaved only); T E , terrestrial) including the main plant life-forms has not, to my knowledge, been attempted. In fact, Esser (1984) stated that such an estimate was not possible with the data available at the time of the Commission of the European Communities Symposium, "Interactions between climate and biosphere," held in Osnabriick in 1983. Esser concluded that NPP partitioning need only be estimated for woody and herbaceous plants of the terrestrial biosphere since the other plant life-forms would be of minor importance in modeling pools and fluxes of C. Our estimate contradicts Esser's assumption in that 9% of terrestrial NPP (DM) was estimated to be associated with macrophyte and nonvascular plant life-forms, the remainder to herbaceous (40%) and woody plants (51%) (Fig. 2.6). By considering each plant life-form separately, it is possible to identify the dominant ecosystems associated with the given plant life-form's annual NPP. Angiosperm woody plants. Ecosystem contribution to angiosperm woody plant NPP was estimated at 63% for forest (greater than half in tropical humid), 22% for savanna (greater than half in seasonal), and 5% for desert (Fig. 2.4A). These figures are unlikely to remain constant through time given the dramatic rates of land-use change. Over the period from 1971 to 1990, the forest area in tropical latitudes was estimated to have decreased at a rate of 15.4xl0 6 hay - 1 (Dixon et al. 1994). Following deforestation, roughly 50% is converted to cultivated land, a small percent to human area and the remainder to degraded lands such as chaparral, maquis, and brushland and desert ecosystems (FAO 1990, cited in Houghton 1994). In contrast to land-use change related to deforestation, global warming under the 2xC02 scenario is predicted to lead to increases in the areal extent of tropical humid forest of between 34 to 58% (Smith and Leemans 1990, cited in Dixon and Turner 1991). However, the annual rate of deforestation (above) is almost two times larger than the predicted increase attributed to global warming. Savanna areas are predicted to increase from 10-43% while desert areas may increase or decrease under a 2xC02 environment (Smith and Leemans 1990, cited in Dixon and Turner 1991). Overall, there should be a net decline in the annual NPP of angiosperm woody plants related to tropical deforestation. Increases may be expected in other ecosystems (savanna and desert), but these will be small in comparison to the losses accompanying deforestation. Gymnosperm woody plants. Ecosystem contribution to gymnosperm woody plant NPP was estimated at 78% for forest (primarily from boreal, temperate, and tropical ecosystem-types), 7% for temperate woodland, and 5% for chaparral, maquis, and brushland (Fig. 2.4B). Total land area associated with boreal forest is predicted to decrease 15 to 44%, with temperate forest 14 to 26%, and with chaparral, maquis, and brushland is predicted to either increase or decrease under a 2xC02 scenario (Smith and Leemans 1990, cited in Dixon and Turner 1991). In their estimate, areal extent of boreal forest was more than two times that of temperate forest. Thus, there may be a net decrease in the annual NPP of gymnosperm woody plants in a 2xCC>2 environment. This implies that future gymnosperm NPP, compared to current NPP levels in boreal forests, will be proportionally greater in temperate forest and perhaps in chaparral, maquis and brushland, which is may increase following deforestation (see above). Herbaceous plants. Ecosystem contribution to herbaceous plant NPP was estimated at 49% for savanna, (nearly equal contributions from the 3 ecosystem-types), 26% for cultivated land (90% from herbaceous reproductive crops), and 18% for temperate grassland (62% moist) (Fig. 2.4C). As stated above, savanna area may increase in a 2xC02 environment. Temperate grassland area may increase or decrease (Smith and Leemans 1990, cited in Dixon and Turner 1991). If grasslands migrate North during the expected global climate change, replacing forests (Apps et al. 1993), their global cover may increase. Whereas, if grasslands are more heavily grazed as pasturelands for a long-term period, they may decrease. Cultivated land is expected to increase to meet the demands of the growing population. In 1990, the establishment of cultivated land was roughly 10x10^ hay - 1 (Houghton 1994). In summary, herbaceous plant contribution to global NPP is expected to increase at the expense of both angiosperm and gymnosperm woody plants, as boreal and tropical forests are replaced with savannas, cultivated lands, and temperate grasslands. Macrophytic plants. Ecosystem contribution to emergent macrophytic plant NPP was estimated at 98% for swamp, marsh, and estuary (mainly saltwater marsh/estuary). Ecosystem contribution to submergent/floating-leaved macrophytic plant NPP was estimated at 99% for swamp, marsh, and estuary (mainly saltwater marsh/estuary). Most of the global macrophyte NPP is thus associated with the saltwater marsh/estuary ecosystem-type which is primarily associated with the submergent/floating-leaved macrophytic life-form. Estimates of changes in the area of this ecosystem are not available. However, a reasonable prediction is a decrease associated with increasing pollution from terrestrial run-off, which contains pollutants. Freshwater swamps, marshes, and lakes, associates with the remaining macrophyte NPP, wil l decrease in area in a 2xC02 environment and with deforestation (see above), since 20% of the areal extent of wetlands is located in tropical humid and boreal ecosystems (Mitsch and Gosselink 1993). Thus, the macrophytic life-form may experience a net decrease in global NPP as the ecosystems associated with its habitat decrease in areal extent. Nonvascular plants. Ecosystem contribution to nonvascular aquatic (non-macrophyte) plant NPP was estimated at 95% for marine (Fig 2.4). Changes in marine land-use are not likely to limit growth of this plant life-form and may even stimulate growth as run-off from terrestrial ecosystems contains growth-limiting inorganic nutrients. Ecosystem contribution to nonvascular aquatic (macrophyte) plant NPP was estimated at 50% for swamp, marsh, and estuary and 34% for algal bed and reef. NPP associated with this plant life-form may decrease due to changes in ecosystem areas as described above for macrophytes. Ecosystem contribution to nonvascular terrestrial plant NPP was estimated at 28% for forest (mainly tropical humid and boreal forest eco-types), 25% for tundra, and 21% for peatland. Since tropical humid and boreal forest land areas are expected to decrease in the future (see above) as are tundra (49-69%, Smith and Leemans 1990, cited in Dixon and Turner 1991) and peatland ecosystems (located in boreal and tundra latitudes), a net decrease in nonvascular terrestrial NPP is expected in the future. Plant organs. Global allocation of NPP (DM) among plant organs was estimated at 30% nonvascular "aboveground parts," 26% fine root, 13% leaf, 12% coarse root/rhizome/tuber (herbaceous); 7% stalk (herbaceous, emergent macrophyte)/petiole (submergent/floating-leafed macrophyte), 5% stem, 4% branch, and 3% reproductive (vascular) (Fig. 2.6). Thus, the least conspicuous organs in the environment, the nonvascular plant "aboveground parts" and the fine and coarse roots of vascular plants, are associated with nearly 70% of the annual NPP. Belowground, structural and water/nutrient-obtaining organs account for 38% of the annual NPP, nearly two-thirds of which is associated with obtaining water and nutrients. Aboveground, structural support organs are associated with 16%, C acquisition and gas-exchange organs with 13%, and reproductive organs which contain the genetic code of future plants with 3% of annual NPP. In vascular plants, it would appear that the function of obtaining water and nutrients is the most C-demanding process of all. This is consistent with the multi-functional role of water in plants as a cell solvent, a medium of inorganic and organic compound transport, a substrate of photosynthesis, a product of mitochondrial respiration, and a cofactor of numerous biochemical reactions. Stem/stalk/petiole and leaf organs appear to place similar demands on the vascular plant C pool in their functions of providing structural integrity, for the former, and a medium in which to carry out photosynthetic carbon fixation and transpiration for the latter. Perhaps C allocation to these two organ functional-types have a close interaction that provides the maximum C economy to aboveground plant organs. Reproductive organs represent a small C-demand in terms of the overall, annual C budget of vascular plants. At the time of organ construction, however, they may be a large drain on the C pool. Two aspects of plant organ NPP were noticeably different in vascular plants. The first noticeable difference is that of NPP partitioning between stems and branches of woody plants. Angiosperm woody plants were estimated to allocate NPP equally between stem and branch organs. Gymnosperms, however, were estimated to allocate 3.5 times more NPP to stem versus branch organs (Fig 2.6). This may be related to the branching structures, gymnosperms being nearly monopodial and angiosperms polypodial. The second notable aspect of vascular plant organ NPP lies in the belowground system. Woody, herbaceous, and aquatic macrophytes were estimated to allocate roughly 50%, 20%, 65% of their NPP to fine root organs and 5%, 25%, 64% to coarse root/rhizome/tuber organs, respectively. The nearly opposite allocation fine root and coarse root allocation patterns between woody and macrophytic plants may be related to the substratum of growth, terra firma versus aquatic, respectively. In the terrestrial environment, greater fine root production could be linked to water and nutrient acquisition. In aquatic environments, water and thus nutrients may be more plentiful thus leading to greater C allocation to structural roots. Equal C allocation between fine and coarse roots/rhizomes of herbaceous plants may be related to the growth habit. Both root types may serve similar C-demanding functions, the rhizomes/coarse roots for creeping along underground to provide physical bases for the annual production of fine roots and stalks and the fine roots for water/nutrient acquisition (Dong and de Kroon 1994). 76 Ratios of belowground net primary production to aboveground net primary production. Based upon the information presented in Fig. 2.6, the global ratio of B N P P : A N P P (DM) was estimated at 0.6. Thus, nearly 40% of the annual NPP occurs belowground. Considering that NPP in the marine ecosystem, which represents 27.3% of global NPP (Table 2.1), occurs solely aboveground in nonvascular plants, the relatively high belowground contribution to global NPP is attributable to the BNPP:ANPP ratio in terrestrial ecosystems, estimated at 1.1. Such a comprehensive estimate of the contribution of belowground and aboveground plant organs to global NPP was, until now, unavailable in the literature. This is not surprising as portions of this estimate were based upon assumptions of ecological processes across ecosystems rather than documented field studies. Terrestrial BNPP:ANPP ratios ranged from 0 (algal bed and reef) to 3.66 (mangal) in aquatic and from 0.06 (polar desert tundra) to 3.82 (sandy hot and dry and desert and semidesert scrub deserts) in terra firma ecosystems (Table 2.8). That the ratios range from 0 to nearly 4 in both aquatic and terra firma systems is consistent with the diverse nature of plant life-forms and their differential importance in ecosystems. Since the ecosystem NPP partitioning values (Fig. 2.6) are representative of the general ecosystem states that result from plant interactions with abiotic and biotic factors, it is difficult to relate our results to any one factor. A potential indicator of the overall ratio of BNPP: ANPP in some ecosystems is the dominant plant life-form. The BNPP:ANPP ratios associated with the global NPP of the dominant plant life-forms were the following: 0, nonvascular; 1.3, woody; 0.9, herbaceous; 3.0, aquatic macrophyte. The average BNPP:ANPP ratio for forest ecosystem-types is estimated at 1.5 which is consistent with the woody plant as the dominant plant life-form. The average BNPP:ANPP ratio for savanna ecosystem-types is estimated at 1.3 which is consistent with a shared life-form dominance by woody and herbaceous plants. The BNPP:ANPP ratio for the lake and stream ecosystem is 0.04 which is consistent with nonvascular plants as the dominant plant-life form. Exceptions to this rule occur in the mangal forest, boreal forest, high and low arctic/alpine tundra, desert, grassland, and herbaceous root/tuber crop cultivated land ecosystems, all of which have higher BNPP:ANPP ratios, and herbaceous reproductive/shoot crops cultivated Table 2.8 Estimated ecosystem ratios of belowground net primary production (BNPP) to aboveground net primary production (ANPP). E C O S Y S T E M ECOSYSTEM-TYPE RATIO OF BNPP:ANPP Forest Tropical humid 1.09 Tropical seasonal 1.09 Mangal 3.66 Temperate evergreen/coniferous 0.84 Temperate deciduous/mixed 0.84 Boreal coniferous (closed) 2.26 Boreal coniferous (open) 2.26 Plantation <10 y 0.96 Plantation 11 y and older-A 0.96 Plantation 11 y and older-G 0.96 Temperate woodland 1.16 Chaparral, maquis, brushland 1.97 Savanna Nonseasonal 1.16 Seasonal 1.16 Hyperseasonal 1.45 Temperate grassland Moist 0.99 Dry 2.45 Tundra Polar desert 0.06 High arctic/alpine 1.67 Low arctic/alpine 1.67 Desert Desert and semidesert scrub 3.82 Extreme (sandy, hot and dry) 3.82 Extreme (sandy, cold and dry) 0.11 Lake and stream 0.04 Swamp, marsh, and estuary Freshwater swamp 0.75 Freshwater marsh 0.51 Saltwater marsh/estuary 1.52 Algal bed and reef 0 Peatland Bog 0.75 Fen 0.26 Cultivated land Herbaceous reproductive crops 0.29 Herbaceous shoot crops 0.29 Herbaceous root/tuber crops 1.64 Woody reproductive crops 0.96 Woody leaf crops 0.96 Human area 0.84 Marine 0 land ecosystems which have lower BNPP:ANPP ratios than the corresponding dominant life-form, woody in mangal and boreal forest, herbaceous in temperate dry grassland and cultivated land, and shared woody and herbaceous in tundra and sandy hot and desert and semidesert deserts. Possible reasons for the greater allocation of NPP to belowground organs in these ecosystems include the following. In the mangal forest, root organs are under greater demand to provide structural support and maintain adequate oxygen in the wave-inundated environment which is reflected in the extensively developed prop root and pneumatophore coarse root designs. In boreal forest, low soil temperatures, large amounts of precipitation as snow, and short summer season lead to accumulations of forest floor litter which remains undecomposed. This may lock up nutrients leading to stimulation of fine root growth. In temperate dry grassland, low mean annual precipitation in combination with grazing may lead to increased BNPP. In herbaceous cultivated lands, altered BNPP:ANPP ratios are related to the herbaceous crop-type which has been selected, either naturally or by genetic manipulation, to produce either more (root/tuber) or less of the desired organ (reproductive/shoot) at harvest. In tundra, increased D M allocation to belowground organs may be related to survival in the severe cold, permafrost and therefore dry climate. In deserts, high root NPP may also be related to levels of precipitation, especially the seasonal BNPP which occurs following a burst of precipitation (Caldwell and Camp 1974). The most widely studied ecosystem in terms of estimating BNPP and one in which estimates of NPP associated with belowground organs are readily available is the forest ecosystem. Even though comprehensive, many of these estimates, especially those prior to 1980, are conservative since they are based upon studies in which the methodologies used to estimate fine root production did not consider annual turnover (mean life-span), C leaching into the rhizosphere, or C diverted to mycorrhizal growth. These processes are critical to more accurately estimating BNPP. Root turnover has been estimated at between 30 to 86% (Fogel 1983) and greater than 100% (Santantonio and Hermann 1985) of the annual fine root NPP of various forest ecosystems. Carbon leaching from herbaceous plant roots has been estimated at 8-15% of annual C fixation (Biondini et al. 1988) and 5-6% over a two week period of C fixation (Shepherd and Davies 1993). Mycorrhizal fungi comprise between 4 to 17% (vesicular-arbuscular mycorrhizae) and 20-39.1% (ectomycorrhizae) of the dry weight of fine roots (Hurley 1989) and may be associated with 14-15% of annual NPP (temperate coniferous forest, Vogt et al. 1982 in Hurley 1989). Even though the studies utilized in our estimate accounted for fine root turnover, only one measured the mycorrhizal contribution to fine root turnover (10% and 18% of conifer fine root turnover in 23- and 180-year old Abies amabilis ecosystems, respectively, Grier et al. 1981). The other studies may have included mycorrhizal production with fine root production; however, this was not mentioned in the methods sections. None of the studies measured C losses through leaching. Thus, even though the BNPP:ANPP ratios presented for ecosystems in this study may reflect more accurately the contributions from fine roots, they do not consider C losses to mycorrhizal fungi or to the rhizosphere. Thus, the BNPP:ANPP ratios estimated for global ecosystems in this estimate may be underestimates. Implications for modeling global plant net primary production. Models of global plant processes are becoming increasingly important as tools to predict the response of plants to global environmental change. Empirical models are based upon data collected from direct studies of plants, thus, our estimates of plant NPP allocation may be compared to those used to model plant functions on a global scale. Currently, most models rely upon the plant NPP estimates of the 1970s which separated terrestrial plant NPP into ecosystems but not into plant life-forms or organs. Two important pieces of information arise from our global estimate which may be applied to improving the model variables of plant NPP in global models. First, the estimate allocates annual NPP to both aboveground and belowground organs based upon direct measurements documented in the literature. Forest BNPP: ANPP ratios, for example, were estimated to range from 0.96-3.66 (Table 2.8). The older BNPP:ANPP ratios, upon which the global NPP estimates were based, were derived indirectly from either the relation of ANPP: aboveground plant biomass to BNPP:belowground plant biomass or by assuming B N P P to be one-third of annual leaf litterfall. This is evident in the current Terrestrial Ecosystem Model (TEM) developed by Raich et al. (1991) which estimated BNPP indirectly from woody root biomass, stem biomass, and stem NPP data (coarse roots) and soil respiration and litterfall data (fine roots). Based upon these relations, BNPP:ANPP ratios in forests were most likely underestimated. Second, the estimate allocated global NPP among the main classes of plant life-forms and plant organs (Table 2.2). As mentioned previously, macrophytes and nonvascular plants may be associated with 9% of the annual terrestrial NPP. Most literature up to the 1970s did not estimate contribution of these plant life-forms and plant organs to the total terrestrial NPP, leading to underestimates. The Raich T E M estimated leaf and branch NPP indirectly from litterfall measurements and made no corrections for losses to decomposition. The model of forest stand growth by Makela and Hari (1986) employed woody gymnosperm partitioning values of 18% to leaf, 24% to branch, 35% to stem, and 23% to root organs. In our estimate, terrestrial NPP associated with woody gymnosperm plants was estimated at 16% leaf, 6% branch, 20% stem, and 56% root organs. The possible improvements from our estimate are the greater allocation to roots, related to fine root production, and the greater ratio of stem:branch in our estimate, 3.5 compared to 1.4. The simulation study of the global C cycle by Goudriaan and Ketner (1984) applied NPP variables for leaf, branch, stem, and roots organs for five ecosystem groups using the ecosystem NPP values of Ajtay et al. (1979). Their estimate is weak for two reasons. First, they clumped tropical forests, forest plantations, nonseasonal savannas, and chaparral, maquis, and brushland into the category "tropical forest," temperate and boreal forests and woodlands into "temperate forest," tundra and semi-desert into one category, and extreme deserts were not included. With this classification, in combination with their "educated guesses" of NPP allocation to the plant organs in these ecosystems, their model was flawed from the beginning. Their estimated BNPP:ANPP ratios ranged from 0.25 to 0.67 of the total NPP of the ecosystem groups, underestimating BNPP. In addition, no attempt was made to estimate plant organ associations with plant life-forms. Thus, their simulated pools of C associated with these plant organs are too general to provide accurate information. Our research could improve the estimate of Goudriaan and Ketner (1984) by providing information concerning the potential C sinks associated with plant life-forms and plant organs. Emanuel et al. (1984) modeled the role of terrestrial ecosystems as potential C stores under scenarios of land-use change using three global NPP classes, nonwoody parts of trees (22 Pg.y-1), woody parts of trees (25 Pg C-y 1 ) , and ground vegetation (15 Pg C-y 1 ) . No distinction between ANPP and BNPP was provided as was no distinction between the macrophytes, herbaceous plants, and nonvascular plants in the ground vegetation. Woody plants were assigned 76% of the terrestrial NPP which is 1.5 times greater than the terrestrial NPP allocation to woody plants in our estimate. Thus, our estimates may also help to improve the Emanuel ecosystem model. Based upon the few studies which allowed us to estimate BNPP:ANPP ratios, it is apparent that the annual net primary productivity values employed by Ajtay et al. (1979) to estimate total ecosystem NPP may be underestimated. The aboveground rate of net primary production in tropical humid forest, for example, was estimated at 2007 g-m^-y 1 (see Methods). Employing the BNPP:ANPP ratio for tropical humid forest (1.09, Table 2.8), the average rate of net primary production is estimated at 4195 g-nr 2-y _ 1. This value is 1.8 times the value of 2300 g-nr^y" 1 presented by Ajtay et al. (1979). The Raich T E M estimated the tropical humid forest rate of NPP at 2040 g-m^-y 1 , similar to that of Ajtay et al (1979) and with similar limitations as noted previously. Similar underestimates may have occurred in other ecosystems. This information suggests that a re-evaluation of terrestrial NPP is necessary to represent accurately the magnitude and nature of plant processes in C cycle models. In the meantime, our estimates may be used as model variables to improve models already in existence, such as the T E M developed by Raich. Estimate Caveats While this estimate of global NPP allocation among ecosystems, plant life-forms, and plant organs has potential to be a powerful tool in providing modelers with missing information required to predict the changes to vegetation under anthropogenic-induced changes in carbon pools and fluxes, there are a few limitations to our estimate. First caveat. The global NPP values presented in Ajtay et al. (1979) are not representative of the current proportions of NPP associated with ecosystems. Since the Ajtay estimate, changes in land-use, the largest being tropical deforestation (Houghton 1994), have altered the relative allocation of NPP among ecosystems. Although our estimate is based upon the outdated Ajtay ecosystem NPP subdivisions, it still provides the relative allocation patterns among plant life-forms and plant organs. Once updated ecosystem NPP values are obtained, our estimates may be applied to these values. Second caveat. The information which served as the basis for plant NPP allocation, the 160 ecological studies, represents plant NPP processes from short-term field measurements. Rarely were any of the studies conducted over a time frame greater than one year. The number of studies applied to estimating NPP allocation per ecosystem also greatly varied from only a few to well over 20 (tropical humid forest). The number of studies utilized is not necessarily related to the quality of the estimate. Tropical humid forest NPP partitioning was associated with the largest number of studies. However, this was due to the nature of the estimate of NPP in this ecosystem. There were no available studies which allocated NPP among all plant life-forms; therefore, many litterfall studies were used to obtain indirect estimates of plant organ NPP. In addition, most studies were not complete in that NPP partitioning information for each plant life-form and plant organ was not available in each ecological study. To overcome this, assumptions for missing plant life-form and/or organ information were based upon either descriptions of the given entity or information from a physiognomically-related ecosystem or ecosystem-type. Thus, error analysis was not possible on a global scale in this estimate. Third caveat. Although many studies were classified as "incomplete" for the purposes of our estimate, in that they did not consider NPP associated with plant litterfall or decomposition, some underestimates could not be avoided. None of the studies accounted for NPP losses to herbivory which may account for a 5% loss of total NPP based upon various estimates in the literature. Studies of herbaceous plants mainly estimated NPP as the average change in biomass over the growing season or less often the peak biomass at the height of the growing season. No corrections were made for NPP lost to litterfall, therefore, total contribution of herbaceous plant NPP to global NPP may be underestimated. Fourth caveat. Perhaps the most significant unknown in this estimate is that of BNPP. An unavoidable error associated with the BNPP estimation is that of C drain to mycorrhizal fungi. Since only one study acknowledged the contribution of mycorrhizal fungi to fine root NPP, the errors associated with this component of NPP are unknown. In order to minimize the errors associated with estimating BNPP, partitioning values were employed only if the values made corrections for mortality and decomposition losses to NPP thoughout the sampling period. Even with this attempt to eliminate error in the BNPP estimate, it was impossible to account for the under- or over-estimates associated with the various methodologies employed by researchers to estimate root turnover (Fogel 1983; Hendricks et al. 1993; Kurz and Kimmins 1987). CHAPTER 3. PATHOGEN-INDUCED ALTERATIONS OF RESPIRATORY C A R B O N FLOW 84 INTRODUCTION Plants respond to stress such as wounding, U V irradiation, and fungal infection by increasing the activity of the OPP, shikimic acid, and phenylpropanoid pathways to synthesize defensive and protective compounds. An important role for respiratory metabolism, including increases in both glycolysis and the OPP pathway, has been demonstrated in the long-term response of a wide variety of plant tissues to wounding (Daly 1976; Kahl 1974). The increase in the activity of the OPP pathway coincides with the lignification associated with wounding in Coleus blumei and Helianthus tuberosus (Pryke and ap Rees 1976) and Pisum sativum (Wong and ap Rees 1971) and provides much of the reductant required for lignification (Pryke and ap Rees 1977). In other types of stress the overall response resembles that seen in wounding but the role of respiration has not been elucidated. Response to pathogenic attack can be studied in vitro using parsley (Petroselinum crispum L.) cell suspension cultures and an elicitor purified from the fungus Phytophthora megasperma f.sp. glycinea, Pmg. Following treatment with Pmg elicitor, transcription and enzyme activity associated with the shikimic acid pathway (Henstrand et al. 1992), phenylpropanoid metabolism (Hahlbrock and Scheel 1989), and the production of defensive furanocoumarins (Hauffe et al. 1986; Kawalleck et al. 1992) increase. Furanocoumarin biosynthesis depletes the substrates of the shikimic acid pathway, the pathway linking respiratory metabolism to phenylpropanoid metabolism (Fig. 1.1), which places a significant demand on primary carbon metabolism to supply E4P and PEP. Demands for other products of primary metabolism, such as ATP and reductant, may be associated with the following events which occur during the initial phases of elicitor recognition by parsley: intracellular acidification (Kneusel et al. 1989), extracellular alkalinization (Scheel et al. 1991), influx of C a 2 + and H + and efflux of CI" and K + (Conrath et al. 1991, Sacks et al. 1993), and Ca2+-dependent phosphorylation/dephosphorylation of proteins (Dietrich et al. 1990; Renelt et al. 1993). Thus, primary carbon metabolism should be activated during parsley elicitation in order to provide the substrates for both the immediate and long-term responses. Research on the response of parsley to the Pmg elicitor has focused on the transduction of the elicitor's signal to the nucleus and the activation of genes coding for enzymes involved in the production of furanocoumarins and other minor phenolics while the role of primary carbon metabolism in the defensive response remains poorly understood. In this study, we have examined the activation of respiratory carbon metabolism in elicited parsley cell suspension cultures. Results are discussed in terms of a role for both the immediate and long-term activation of glycolysis and the OPP pathway in the defensive response. 86 METHODS Cell culture and elicitor Cell suspension cultures of parsley (Petroselinum crispum L.) (Schmelzer et al. 1985) were grown in the dark (150 rpm, 25°C) in 250 mL or 1 L Erlenmeyer flasks as previously described (Ragg et al. 1981) and were subcultured in fresh modified B5 medium (Hahlbrock 1975) every seven days. Experiments were performed on six day old cultures that had reached a density of 120-140 mg m L - 1 fresh weight as determined after vacuum filtration using Whatman filter paper (#1) for 10 seconds. The Pmg elicitor, a crude cell wall extract of the fungus Phytophthora megasperma f. sp. glycinea, isolated as described by Ayers et al. (1976), was kindly donated by Dr. Carl J. Douglas (University of British Columbia), and aliquots of a stock solution (5 mg m L - 1 ) were used in all elicitation experiments to give a final concentration of 30 |J,g m L - 1 . In control experiments, Pmg solution was replaced by an equivalent volume of distilled, deionized water (ddH20). Gas exchange experiments A l l experiments were performed in the dark at 25°C in a sterile 160 mL water-jacketed chamber attached to an infrared gas analyzer (IRGA, The Analytical Development Co. Ltd., England) in an open gas exchange system. Sterile C02-free air was channeled from the IRGA to the bottom of the chamber at 200 m L m i n - 1 and then returned to the IRGA. The IRGA, calibrated with 90 | i l L _ 1 CO2 in compressed air, was used to monitor the CO2 concentration in the air returning from the cell suspension. Cells (40-50 mL) were transferred aseptically to the chamber through a sealed sampling port at the base of the chamber using 60 mL syringes capped with 18.5G (wide-bore) needles and buffered with 25 m M Mes, pH 5.7. Cell samples were removed with syringes capped with 18.5G (wide-bore) needles through the sampling port. Cells were allowed to equilibrate for 30 min prior to treatment to obtain a steady rate of C O 2 evolution. Pmg or ddH20 was added, and the responses were monitored for 100 min. Both elicitation and control experiments were performed in triplicate. Three 1 mL samples were taken at the end of the experiment for protein determination. Rates of CO2 evolution were expressed as iimol CO2 mg - 1 protein tr 1 . Metabolite experiments Metabolite experiments were performed in triplicate under the same conditions described above for gas exchange experiments except 1 mL metabolite samples were withdrawn through the sampling port at points throughout the experiment and three 1 mL samples were removed prior to the experiment for protein determination. Samples were removed, plunged into 167 ul of 70% H C I O 4 , giving a final concentration of 10% (v/v), and rapidly frozen in liquid N2. The time course of metabolite sampling included 10 min prior to treatment and 100 min following treatment. Metabolite samples were thawed, centrifuged (15g x 103, 4 min), and the supernatant was neutralized with 5N K O H / 1 M triethanolamine. The samples were again centrifuged, and the supernatants were brought up to 1.5 mL with ddH20 and stored in liquid N2 until analyzed. Metabolites were measured in enzymatic assays coupled to the reduction/oxidation of pyridine nucleotides using a dual-wavelength U V - V I S spectralline-photometer (ZFP-22, Sigma Instruments, FRG). Glucose-6-phosphate (Glc-6-P), fructose-6-phosphate (Fru-6-P), fructose-1,6-bisphosphate (Fru-l,6-bisP), and 6-phosphogluconate (6-PG) assays were performed as previously described (Quick et al. 1989; Wirtz et al. 1980). Metabolite recoveries of 88.0% (Glc-6-P), 82.4% (Fru-6-P), 95.4% (Fru-l,6-bisP), and 81.2% (6-PG) were determined by extracting known amounts of metabolites along with the samples. Corrections for any changes in protein concentration during the experiment were accounted for in the calculations. C(yiCi ratios: analysis of14CO2 evolution Parsley cells were prepared and equilibrated as previously detailed (see above). Aliquots of cells (5 mL) were removed immediately before and 60 min following Pmg or ddH^O addition. Samples were placed in 25 mL Warburg flasks containing 1.0 mCi (5.0 fiL) of either D - [1- 1 4 C] or D-[6- 1 4 C] glucose (60 mCi-mmoW) with a center well containing 1 mL of the basic CO2 trap, B-phenylethylamine. The flasks were incubated for 5 min with continuous shaking and all reactions were stopped by the addition of 0.72 mL 70% H C I O 4 . Flasks were shaken an additional 24 hours to ensure complete CO2 release from the medium and effective trapping of C O 2 . The center well contents were removed and counted in American Chemical Society scintillation cocktail using a Beckman LS 60001C scintillation counter and the results used to calculate the C6:Ci ratios of 14CC>2 evolution. Other methods Protein was measured utilizing the Bradford Assay (Bradford 1976) with gamma-globulin as the standard. The stimulation of furanocoumarin synthesis was routinely confirmed by monitoring U V fluorescence of the furanocoumarins 24 hours post-elicitation (Kombrink and Hahlbrock 1986). RESULTS Respiratory CO2 evolution The rate of CO2 evolution increased 61% from 2.46 to 3.96 jxmol mg - 1 protein h _ 1 (Fig. 3. IB) within 20 min of elicitor treatment. The rate increased more slowly over the next 40 min, reaching a steady rate of 4.42 |j,mol mg - 1 protein tr 1 which represents an overall increase of 80%. This biphasic pattern of CO2 evolution was observed in all experiments. The enhanced rate of C O 2 evolution was maintained for the duration of the experiment. The rate of C O 2 evolution in the control experiments rose from 2.19 to 2.37 |imol m g - 1 protein h r 1 after 100 min, an insignificant increase of 8% (Fig. 3.1 A), indicating that transfer of cells from the growing conditions to an aerated environment did not significantly affect the rate of dark respiratory CO2 evolution over the duration of the experiment. Metabolites Elicitation had no effect on the levels of fructose-6-phosphate (Fru-6-P), the substrate of phosphofructokinase (PFK), throughout the experiment (Fig. 3.2A). The levels of fructose-1,6-bisphosphate (Fru-l,6-bisP), the product of PFK, dropped within the first 6 min of elicitation and then rose to a level 2 times the control (Fig 3.2B). Likewise, the ratio of Fru-l,6-bisP:Fru-6-P began to increase within 7 min following elicitation (Fig. 3.2C) and reached a maximum 2 times that of the control within 20 min. The rapid and sustained increase in the Fru-l,6-bisP:Fru-6-P ratio is consistent with the activation of P F K for the duration of the experiment. The levels of glucose-6-phosphate (Glc-6-P), the substrate of glucose-6-phosphate dehydrogenase (G6PDH), decreased within 12 min following elicitor treatment, returning to control levels thereafter (Fig. 3.3A). A corresponding increase in the levels of 6-phosphogluconate (6-PG), the product of G6PDH, was observed within 4 min (Fig. 3.3B), reaching a level 7 fold higher than the control within 20-30 min of elicitation. The ratio of 6-PG:Glc-6-P began to increase within 4 min of elicitation and reached maximum levels, 7 times 90 Figure 3.1 Representative traces of the changes in the rate of CO2 evolution in cell suspensions of Petroselinum crispum treated with water (A) or Pmg elicitor (30 ligmL" 1) (B). Traces are from two individual experiments and do not corresond to the average values for each treatment obtained from three separate experiments. The increase in the rate of CO2 evolution in (A) is insignificant at 100 min following treatment (paired-sample t test, p>0.05) and in (B) is significant at 20 and 60 min following treatment (paired-sample t test, p<0.001). 91 5.0 4.0 3.0 2-0 [• Q l I CC Q. 1 t= V 0 "I § | 4 0 O - S 3.0 U 2.0 1.0 1.4 0.2 g 1.0 I-< 0.6 I--20 A. Fru-6-P B. Fru-1,6-bisP _L C. Fru-1,6-bisP:Fru-6-P _L 20 40 60 80 100 120 TIME (min) Figure 3.2 The intracellular levels of Fru-6-P (A) and Fru-l,6-bisP (B) and the ratio of Fru-l,6-bisP:Fru-6-P (C) in cell suspensions of Petroselinum crispum treated with Pmg elicitor (30 iig-mL"1) (O) and water (•). Symbols represent the mean for 3 separate experiments. Arrow indicates time of treatment. The error bars on the last time point represent the average SE of the three experiments for each treatment. 92 9 1 cz CD •I—» O < DC Q-h- v z co LU £ O -= ° i o 3 o h-< rr 13 12 -11 -10 -9 \-8 7 0.8 -0.6 -0.4 -0.2 0 0.08 0.06 0.04 0.02 0 A. Glc-6-P B. 6-PG C. 6-PG:Glc-6-P -20 0 T 20 40 60 80 100 120 TIME (min) Figure 3.3 The intracellular levels of Glc-6-P (A) and 6-PG (B) and the ratio of 6-PG:Glc-6-P (C) in cell suspensions of Petroselinum crispum treated with Pmg elicitor (30 j l g m L - 1 ) (O) and water (•). Symbols represent the mean for 3 separate experiments. Arrow indicates time of treatment. The error bars on the last time point represent the average SE of the three experiments for each treatment. greater than those of the control, within 20-30 min (Fig. 3.3C). The increase in the 6-PG:Glc-6-P ratio upon elicitation indicates the immediate and sustained activation of G6PDH. C(,.Ci ratios of14CO2 evolution The C6:Ci ratios determined prior to and 60 min following treatment are presented in Table 3.1. Prior to elicitation, there was no significant difference between control and treatment with average ratios of 0.46 and 0.43, respectively. After 60 min, the control ratio was not significantly affected with a value of 0.44 while the treatment ratio dropped to 0.33, a 23% decrease. 94 Table 3.1. Effect of Pmg elicitor (30 jLtg -mL" 1 ) on the C6:Ci ratios of 1 4 C 0 2 evolution from cell suspensions of Petroselinum crispum. (mean ± SD). Numbers in parentheses represent the number of times the experiments were duplicated. Values with different letters are significantly different at the 99% confidence interval using the Kruskal-Wallis test for multiple sample comparisons. TIME (min) T R E A T M E N T 0 60 PERCENT C H A N G E Control «0.46 ± 0.07 (7) a0.44 ± 0.05 (11) -4 Pmg «0.43 ± 0.05 (7) ^0.33 ± 0.05 (8) -23 DISCUSSION Immediate events following elicitor treatment In parsley cells elicited with Pmg there is an immediate increase in the rate of CO2 evolution (Fig. 3.1). The increase in the ratios of Fru-l,6-bisP:Fru-6-P (Fig. 3.2) and 6-PG:Glc-6-P (Fig. 3.3) within minutes of elicitation indicate the activation of two respiratory enzymes, PFK and G6PDH, respectively. The timing of these changes supports a role for the activation of primary metabolism in the elicitor recognition and signal transduction processes that activate secondary metabolism. During this time frame there are dramatic changes in ion fluxes across the plasma membrane and changes in the phosphorylation status of proteins. Activation of primary metabolism under these conditions would serve to provide ATP and reducing power to meet the increased cellular energy demands arising from elicitor-induced stress. P F K and G6PDH are key regulatory enzymes in glycolysis and the OPP pathway, respectively, and are finely regulated by a number of factors (Dennis and Miernyk 1982; Copeland and Turner 1987, Plaxton 1990). Immediate changes in intracellular pH could be a key factor in the activation of PFK. Small pH changes can lead to large changes in PFK activity in some plant systems (Kelly and Turner 1969; Botha et al. 1988) As previously demonstrated by Kneusel et al. (1989), the cytoplasmic pH of aerated parsley cells decreases as much as 0.25 pH unit following treatment with Pmg. The timing of this pH drop corresponds to the alkalinization of unbuffered external medium (data not shown) consistent with pH playing a role in the activation of PFK in elicited parsley cells. P F K is also under negative regulatory control by PEP and ATP in higher plants (Dennis arid Greyson 1987). The immediate, transient drop in the levels of Fru-l,6-bisP (Fig. 3.2B) and triose-phosphate (data not shown) may indicate activation of another rate-limiting glycolytic enzyme, pyruvate kinase, which regulates the flow of PEP to the T C A cycle (Fig. 1.1). Alternatively, there could be an immediate drain on PEP pools for increased secondary metabolism, committing carbon to the shikimic acid pathway. The pools of A T P may be utilized during elicitor recognition and ion transport at the plasma membrane. A drop in the levels of either of these metabolites could result in the activation of PFK. Unfortunately, poor yields of PEP and pyruvate in this study prevented reliable measurements of changes in these metabolites, while ATP measurements were inconclusive. In response to elicitation, the 6-PG:Glc-6-P ratio increases rapidly indicating the immediate activation of G6PDH (Fig. 3.3). In many plant systems the principle activator of G6PDH is a low NADPH:NADP+ ratio. Elicitation induces an immediate extracellular oxidative burst which involves the extracellular production of O2 ~ and subsequent transient formation of H2O2 within 1-4 min of elicitation in many plant-pathogen interactions (Doke 1983; Apostol et al. 1989) including the interaction of parsley cells with the Pmg fungal elicitor (Dierk Scheel, personal communication). The production of H2O2 from 0 2 ~ is thought to involve a membrane bound NADPH-utilizing oxidase which transfers 2 H+ and 2 e - from intracellular N A D P H to O2" (Bowler et al. 1992). The immediate production of H2O2 upon elicitation could cause the levels of N A D P H to drop, decreasing the ratio of N A D P H : N A D P + , thereby activating G6PDH. The immediate demand for N A D P H would be consistent with the rapid activation of G6PDH prior to the expected long-term demand for OPP pathway carbon skeletons from the shikimic acid pathway. Long-term events following elicitation The activation of PFK and G6PDH during elicitation indicates that both glycolysis and the OPP pathway contribute to the immediate respiratory enhancement in elicited parsley cells. In the long-term, an increase in secondary metabolism to supply the substrates for furanocoumarin synthesis has been associated with elicitation by Pmg. Several observations support a continued involvement of both glycolysis and the OPP pathway in this long-term response. The elevated rate of CO2 evolution (Fig. 3.1) and the increased product:substrate ratios for G6PDH and PFK after elicitation (Figs. 3.2 and 3.3) are maintained for the duration of the experiment. The G6PDH and 6-phosphogluconate dehydrogenase reactions of the OPP pathway supply the bulk of the reductant required in the shikimic acid pathway and several steps in the formation of furanocoumarins (Fig. 1.1). The increased oxidation of N A D P H via increased secondary metabolism would maintain a low N A D P H : N A D P + ratio, thus supporting the activation of G6PDH and consequently OPP pathway activity. Furanocoumarin production also requires ATP and PEP, both inhibitors of PFK. A demand for these metabolites should keep them at low levels in the long-term, thus sustaining the activation of PFK and hence glycolytic activity. The decrease in the C6:Ci ratio of 1 4 C 0 2 evolution after 60 min (Table 3.1) indicates that the OPP pathway is responsible for some portion of the 80% enhancement of the rate of C O 2 evolution in the presence of elicitor. The decrease in the C6:Ci ratio can be explained by the fact that the CQ carbon of glucose is not oxidized when metabolized through the OPP pathway. In contrast, both the C6 and C] carbons are oxidized when metabolized through the T C A cycle via glycolysis (Fig. 1.1). If the C 6 : C i ratio remained unchanged then we could estimate that the relative contributions of both pathways were unaffected by elicitation even though the total magnitude may have increased. A C6:C[ ratio of 0.24 would be expected if the enhanced respiration (80%) were due solely to the OPP pathway. The observed ratio after 60 min, 0.33 (Table 3.1), is greater than the theoretical value (0.24) but lower then the initial or control value (0.43). This suggests that the relative contribution of the OPP pathway to carbon oxidation has increased to a greater extent than that of glycolysis following elicitation. The incomplete oxidation of triose-phosphate would also contribute to the reduction of the C6:Ci ratio as PEP is diverted away from the T C A cycle to the shikimic acid pathway (Fig. 1.1). Demand for PEP in the DAHP synthase and 5-eno/pyruvylshikimate-3-phosphate synthase reactions of the shikimic acid pathway, leading to the production of furanocoumarins, could lead to the incomplete oxidation of substantial amounts of PEP. The DAHP synthase reaction, which combines one molecule each of PEP and E4P, is the first step committing carbon flow to the shikimic acid pathway, and increased phenylpropanoid metabolism would create a demand for both of these respiratory products. Theoretically, 2 moles of Glc-6-P metabolized through the OPP pathway generate 1 mole each of E4P and Fru-6-P. The Fru-6-P can be metabolized through glycolysis to yield 2 moles of PEP. This PEP can then be used in the D A H P and 5-eno/pyruvylshikimate-3-phosphate synthase reactions which lead to the formation of 1 mole of phenylalanine. In terms of a carbon budget, the two moles of Glc-6-P which enter the OPP pathway represent precisely the number of carbon skeletons required to satisfy the needs of phenylpropanoid metabolism (Fig. 1.1). Evidence linking the increased OPP pathway activity 60 min after elicitation of parsley cells to the production of furanocoumarins includes increased transcription of D A H P synthase (Henstrand et al. 1992) as well as phenylalanine ammonia lyase and 4-coumarate:CoA ligase (Somssich et al. 1986), enzymes of the phenylpropanoid pathway. Increases in the enzyme activities generally have been reported to occur 2 to 3 hours following elicitation (McCue and Conn 1989, Kuhn et al. 1984); however, these activity measurements often represented the first time points taken following elicitation so any earlier activation would have been missed. We attempted to demonstrate a direct link between increased OPP pathway activity and the production of furanocoumarins by using [U- 1 4 C] phenylalanine and [6- 1 4 C] glucose to label newly synthesized furanocoumarins. Unfortunately, the elicitor interfered with the uptake of the substrates by parsley cells (data not shown) which is consistent with the membrane changes reported after elicitation and with the elicitor-induced inhibition of asparagine uptake demonstrated in parsley cells (Strasser and Matern 1986). 99 CHAPTER 4: CONCLUSIONS Global Carbon Allocation Lignin and holocellulose production. Annual net flow of photosynthetically fixed C to lignin and holocellulose synthesis was estimated at 48% of global NPP. Thus, nearly half of net C assimilation is allocated to plant cell wall material. The potential fate of this C in global ecosystems is dependent upon the given ecosystem and the relative content of lignocellulose in the plant organ. Therefore, lignocellulose production in terrestrial ecosystems was estimated to be a potential annual sink of 17.7 Pg C y 1 while holocellulose production in the marine ecosystem was estimated to be a small potential annual sink of 0.3 Pg C - y 1 . Considering land-use changes, including tropical deforestation and savanna fires, a more representative value is 7.9 Pg C - y 1 . This value is greater than that predicted in current publications for terrestrial vegetation as it considers the potential C sinks in cultivated land, grassland, and swamp, marsh, and estuary ecosystems in addition to Northern Hemisphere forests which have recently been implicated as the primary terrestrial C sink. Ecosystems in which high C-sink potential woody stem, branch, and coarse root organs may be a lignocellulose C sink include tropical forests, Northern Hemisphere forests (boreal and temperate), and savannas, for medium C-sink potential woody leaf organs include tropical and Northern Hemisphere forests, for medium C-sink potential woody fine roots include tropical and Northern Hemisphere forests and savannas, for medium C-sink potential herbaceous leaf and stalk organs include nonseasonal savannas and temperate grassland, for medium C-sink potential herbaceous fine root and coarse root/rhizomes include savannas, cultivated land, and temperate grasslands, for medium C-sink potential macrophytic fine root coarse roots/rhizome organs include swamp, marsh, and estuary, and for nonvascular aquatic (non-macrophyte) include marine. Associated with the annual net synthesis of lignin, l x l O 1 4 moles of N are estimated to be reassimilated in the prephenate-arogenate junction, the metabolic link between primary and secondary metabolism that leads to the production of lignin in vascular plants. This implies a 100 tight regulation of N mobilization to sites of active lignin synthesis. As N is often a limiting nutrient to plant growth in terrestrial and marine ecosystems, this may be a pathway of N conservation in plants and requires further investigation. Production of plant life-forms and plant organs. Estimation of global NPP associated with plant life-forms revealed a similar role for both herbaceous nonvascular aquatic (non-macrophyte) plants. Secondary to these were angiosperm and gymnosperm woody plants, and of minor importance were emergent and submergent/floating-leaved macrophytes, nonvascular terrestrial, and nonvascular aquatic (macrophyte) plants. Changes in ecosystem areal cover either from land-use change or global warming under a 2xCC>2 environment may result in a net decrease in the annual NPP of angiosperm woody plants (tropical deforestation), gymnosperm woody plants (boreal forest-global warming decrease), macrophytes (tropical and boreal latitudes), and nonvascular plants (tropical and boreal latitudes), but a net increase in the annual NPP of herbaceous plants (temperate forest, savannas, cultivated land-global warming increases). Estimates of annual NPP associated with global plant life-forms were further subdivided to plant organs, the most C-demanding being fine root, followed by leaf and coarse root/rhizome organs, followed by stalk/petiole, stem, branch, and reproductive organs. This supports a role for water acquisition as the most C-demanding part of plant growth, secondary being C acquisition, and tertiary being structural support and reproduction. Implications for models. Utilizing these improved estimates of plant NPP allocation among ecosystems, plant life-forms, and plant organs, models that require NPP values as parameters may be upgraded. Current models assume little importance of plant life-forms other than woody and herbaceous. In addition, model parameters of NPP associated with woody and herbaceous plant organs were not consistent with this global estimate. My estimate also suggests potential underestimates of BNPP to total NPP of various terrestrial ecosystems including the dominant forest ecosystem. The estimates of lignin and holocellulose production may also be 101 incorporated into models designed to predict changes in pools and fluxes of C under anthropogenically-induced global warming and land-use change scenarios. In conclusion, although there are a number of errors associated with this estimate of global NPP allocation, it brings to light the knowledge gaps present in current theories of NPP. 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