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Photosynthesis and respiration in five species of benthic foraminifera that host algal symbionts

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Title:
Photosynthesis and respiration in five species of benthic foraminifera that host algal symbionts
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English
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Walker, Robert A., 1965-
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University of South Florida
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Subjects / Keywords:
Archaias
Cyclorbiculina
Amphistegina
metabolism
irradiance
Dissertations, Academic -- Marine Science -- Masters -- USF   ( lcsh )
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government publication (state, provincial, terriorial, dependent)   ( marcgt )
bibliography   ( marcgt )
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non-fiction   ( marcgt )

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Summary:
ABSTRACT: Oxygen production and consumption were measured in five species of benthic foraminifers using a "Clark-type" oxygen electrode. Net photosynthesis and respiration were calculated and normalized to both &#956g Chl a and mm² upper surface area for the chlorophyte-bearing soritid foraminifers, Archaias angulatus and Cyclorbiculina compressa, and the diatom-bearing amphisteginids, Amphistegina gibbosa, A. lessonii and A. radiata. Photosynthesis/Irradiance curves were generated by fitting data to the hyperbolic tangent equation P = Pmax tanh (&#945 I/ Pmax). Derived photosynthetic parameters, Pmax, &#945, Ik were found to correspond to the general responses of the endosymbiont taxa. Chlorophyll concentration was found to be significantly lower in Cyclorbiculina compressa than in the other four species. Maximum O₂ production (Pmax) when normalized to Chl a was 3-4 times higher in soritid species than in amphisteginids. Photosynthetic efficiency (&#945) was significantly higher in Amphistegina gibbosa and A. lessonii than in the soritids. Mean Ik, which indicates approaching light saturation, was 13 and 26 &#956mol photon m ⁻2sec⁻1 respectively for A. gibbosa and A. lessonii compared with 95 and 119 &#956mol photon m⁻2sec⁻1 respectively for Archaias and Cyclorbiculina. Calculated P/I data were to variable for Amphistegina radiata to estimate reliable &#945 and Ik values. Factorial metabolic scope, which indicates potential for activity was only 2-6 for amphisteginids versus 9-16 for soritids. Annual primary production was estimated to be 285 mmoles O₂ m⁻2 of habitat for A. angulatus, 9.3 mmoles O₂ m⁻2 of habitat for C. compressa and 15.3 mmoles O₂ m⁻2 of habitat for Amphistegina lessonii. Pmax values for Amphistegina gibbosa fluctuated at the compensation point and did not indicate significant oxygen production. Pmax values for Amphistegina radiata failed to reach the compensation point and net oxygen production was not recorded.
Thesis:
Thesis (M.S.)--University of South Florida, 2004.
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Includes bibliographical references.
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by Robert A. Walker.
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Document formatted into pages; contains 112 pages.

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Photosynthesis and Respiration in Five Species of Benthic Foraminifera that Host Algal Symbionts by Robert A. Walker A thesis submitted in partial fulfillment of the requirements for the degree of Master of Science College of Marine Science University of South Florida Major Professor: Pamela Hallock Muller Ph.D. Gabriel Vargo Ph.D. Joseph Torres Ph.D. Date of Approval: May 27, 2004 Keywords: Archaias Cyclorbiculina Amphistegina metabolism, irradiance Copyright 2004, Robert A. Walker

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Acknowledgements Aspects of this research were supported by grants from the National Science Foundation OCE-92-3278 and CHE-0221834, National Oceanic and Atmospheric Administrations National Undersea Research Progr am Subcontracts No. 9120, 9204.4, 9221, 9322, 9515, 9609, 9703.66 and 9922, by the U.S. Environmental Protection Agency-ORD-STARGAD-R825869; and by South Carolina Sea Gr ant Consortium Project No. R/MB-2.

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i Table of Contents List of Tables iii List of Figures vi Abstract viii Introduction 1 Taxa Studied 2 Estimating Photosynthesis and Respiration 6 Thesis Objectives 9 Methods 11 Collection and Storage of Specimens 11 Photosynthesis and Respiration Trials 14 Chlorophyll Extraction 17 Data Analysis 17 Results 21 Archaias angulatus 21 Cyclorbiculina compressa 26 Amphistegina gibbosa 32 Amphistegina lessonii 37 Amphistegina radiata 40 Discussion 45 Photosynthesis in Symbiotic relationships 47 Photosynthetic Efficiency 52 Irradiance Measures 55 Photosynthesis and Irradiance Raw Data 59 Photoinhibition 62 Respiration 63 Symbiont Bearing Foramini fers as Primary Producers 65 Recommendations for Future Research 68 Conclusion 70 References 72

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ii Appendices 79 Appendix A 80 Appendix B 83 Appendix C 87 Appendix D 93 Appendix E 97

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iii List of Tables Table 1. Dates, locations, and dept hs of collection of foraminifers used in this study. 12 Table 2. Physical parameters of Archaias angulatus 21 Table 3. Derived photosynthetic pa rameters normalized to Chl a for Archaias angulatus 24 Table 4. Derived photosynthetic pa rameters normalized to surface area for Archaias angulatus 24 Table 5. Physical parameters of Cyclorbiculina compressa 27 Table 6. Derived photosynthetic pa rameters normalized to Chl a for Cyclorbiculina compressa 30 Table 7. Derived photosynthetic pa rameters normalized to surface area for Cyclorbiculina compressa 30 Table 8. Physical parameters of Amphistegina gibbosa 33 Table 9. Derived photosynthetic pa rameters normalized to Chl a for Amphistegina gibbosa 34 Table 10. Derived photosynthetic pa rameters normalized to surface area for Amphistegina gibbosa 35 Table 11. Physical parameters of Amphistegina lessonii 37 Table 12. Derived photosynthetic pa rameters normalized to Chl a for Amphistegina lessonii 39 Table 13. Derived photosynthetic pa rameters normalized to surface area for Amphistegina lessonii 39 Table 14. Physical parameters of Amphistegina radiata 41

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iv Table 15. Derived photosynthetic pa rameters normalized to Chl a for Amphistegina radiata 43 Table 16. Derived photosynthetic pa rameters normalized to surface area for Amphistegina radiata 43 Table 17. Annual primary production of A. angulatus C. compressa and Amphistegina lessonii 67 Table A-1. Physical characteristics of Archaias angulatus 78 Table A-2. Chlorophyll a extraction Archaias angulatus 78 Table A-3. Oxygen Production at e xperimental light intensities Archaias angulatus (nmoles hr-1 ug Chl a-1 ) 79 Table A-4. Oxygen Production at e xperimental light intensities Archaias angulatus (nmoles hr-1 mm-2) 80 Table A-5. Metabolic scope and factorial metabolic scope for Archaias angulatus 81 Table A-6. Derived parameters from Photosynthesis/Irradiance curves for Archaias angulatus 81 Table B-1. Physical characteristics of Cyclorbiculina compressa 82 Table B-2. Chlorophyll a extraction Cyclorbiculina compressa 82 Table B-3. Oxygen Production at e xperimental light intensities Cyclorbiculina compressa (nmoles hr-1 ug Chl a-1 ) 83 Table B-4. Oxygen Production at e xperimental light intensities Cyclorbiculina compressa (nmoles hr-1 mm-2) 84 Table B-5. Metabolic scope and factorial metabolic scope for Cyclorbiculina compressa 85 Table B-6. Derived parameters from Photosynthesis/Irradiance curves for Cyclorbiculina compressa 85 Table C-1. Physical characteristics of Amphistegina gibbosa 86 Table C-2. Chlorophyll a extraction Amphistegina gibbosa 87

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v Table C-3. Oxygen Production at e xperimental light intensities Amphistegina gibbosa (nmoles hr-1 ug Chl a-1 ) 89 Table C-4. Oxygen Production at e xperimental light intensities Amphistegina gibbosa (nmoles hr-1 mm-2) 90 Table C-5. Metabolic scope and factorial metabolic scope for Amphistegina gibbosa 91 Table C-6. Derived parameters from Photosynthesis/Irradiance curves for Amphistegina gibbosa 91 Table D-1. Physical characteristics of Amphistegina lessonii 92 Table D-2. Chlorophyll a extraction Amphistegina lessonii 92 Table D-3. Oxygen Production at e xperimental light intensities Amphistegina lessonii (nmoles hr-1 ug Chl a-1 ) 94 Table D-4. Oxygen Production at e xperimental light intensities Amphistegina lessonii (nmoles hr-1 mm-2) 94 Table D-5. Metabolic scope and factorial metabolic scope for Amphistegina lessonii 95 Table D-6. Derived parameters from Photosynthesis/Irradiance curves for Amphistegina lessonii 95 Table E-1. Physical characteristics of Amphistegina radiata 96 Table E-2. Chlorophyll a extraction Amphistegina radiata 96 Table E-3. Oxygen Production at e xperimental light intensities Amphistegina radiata (nmoles hr-1 ug Chl a-1 ) 97 Table E-4. Oxygen Production at e xperimental light intensities Amphistegina radiata (nmoles hr-1 mm-2) 98 Table E-5. Metabolic scope and factorial metabolic scope for Amphistegina radiata 99 Table E-6. Derived parameters from Photosynthesis/Irradiance curves for Amphistegina radiata 99

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vi List of Figures Figure 1. Photographs of all species studied 3 Figure 2. Collection sites of Archaias angulatus (KML), Cyclorbiculina compressa (Conch Reef) and Amphistegina gibbosa (Tennessee Reef). 13 Figure 3. Location of Ambitle Island, Papua New Guinea 13 Figure 4. Sample photosynthesi s/irradiance curve showing P max , and I k parameters 18 Figure 5. Correlation of mass (a ) and surface area (b) to Chl a extracted from individual Archaias angulatus 22 Figure 6. Photosynthesis vs Irradiance normalized to Chl a ( Archaias angulatus) 25 Figure 7. Photosynthesis vs Irradi ance normalized to surface area ( Archaias angulatus ) 26 Figure 8. Correlation of Mass and Surface Area to Chl a extracted 27 Figure 9. Mean Oxygen production/consumption normalized to Chl a showing initial increase in oxygen consumption at lowest light intensity in Cyclorbiculina compressa 28 Figure 10. Mean Oxygen production/consumption normalized to surface area showing initial increase in oxygen consumption at lowest light intensity in Cyclorbiculina compressa. 29 Figure 11. Photosynthesis vs irradiance normalized to Chl a for Cyclorbiculina compressa 31 Figure 12. Photosynthesis vs irradi ance normalized to surface area for Cyclorbiculina compressa 32

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vii Figure 13. Correlation of mass (a ) and surface area (b) to Chl a extracted for Amphistegina gibbossa 33 Figure 14. Photosynthesis vs irradiance normalized to Chl a for the amphisteginid species 35 Figure 15. Photosynthesis vs irradian ce normalized to surface area for the Amphisteginid species 36 Figure 16. Correlation of mass (a ) and surface area (b) to Chl a extracted for A. lessonii 38 Figure 17. Correlation of mass (a ) and surface area (b) to Chl a extracted for A. radiata 41 Figure 18. Photosynthesis vs irradiance normalized to Chl a for all species 48 Figure 19. Photosynthesis vs irradian ce normalized to surface area for all species 49 Figure 20. Maximum photosynthesis (P max ) normalized to Chl a 50 Figure 21. Maximum photosynthesis (P max ) normalized to surface area 51 Figure 22. Chlorophyll a concentration for each species 51 Figure 23. Alpha values for all species normalized to Chl a 54 Figure 24. Alpha values for all speci es normalized to surface area 54 Figure 25. Irradiance values (I k ) at maximum photosynthesis for all species 56 Figure 26. Factorial scope (ratio of post-trial respiration rate to pretrial respiration rate) for all species 64

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viii Photosynthesis and Respiration in Five Species of Benthic Foraminifera that Host Algal Symbionts. Robert A. Walker ABSTRACT Oxygen production and consumption were measured in five species of benthic foraminifers using a Clark-type oxygen elec trode. Net photosynthe sis and respiration were calculated and normalized to both g Chl a and mm2 upper surface area for the chlorophyte-bearing soritid foraminifers, Archaias angulatus and Cyclorbiculina compressa and the diatom-bearing amphisteginids, Amphistegina gibbosa A. lessonii and A. radiata. Photosynthesis/Irradiance curves we re generated by fitting data to the hyperbolic tangent equation P = P max tanh ( I/ P max ). Derived photosynthetic parameters, P max , I k were found to correspond to the general responses of the endosymbiont taxa. Chlorophyll concentration was found to be significantly lower in Cyclorbiculina compressa than in the other four species. Maximum O 2 production (P max ) when normalized to Chl a was 3-4 times higher in soritid sp ecies than in amphisteginids. Photosynthetic efficiency ( ) was significantly higher in Amphistegina gibbosa and A. lessonii than in the so ritids. Mean I k which indicates approaching light saturation, was 13 and 26 mol photon m-2sec-1 respectively for A. gibbosa and A. lessonii compared

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ix with 95 and 119 mol photon m-2sec-1 respectively for Archaias and Cyclorbiculina Calculated P/I data were to variable for Amphistegina radiata to estimate reliable and I k values. Factorial metabolic scope, which in dicates potential for ac tivity was only 2-6 for amphisteginids versus 9-16 for soritids. Annual primary production was estimated to be 285 mmoles O 2 m-2 of habitat for A. angulatus 9.3 mmoles O 2 m-2 of habitat for C. compressa and 15.3 mmoles O 2 m-2 of habitat for Amphistegina lessonii P max values for Amphistegina gibbosa fluctuated at the compensa tion point and did not indicate significant oxygen production. P max values for Amphistegina radiata failed to reach the compensation point and net oxygen production was not recorded.

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1 Introduction Larger benthic foraminifers are so abunda nt in many reef environments that they have been called living sands by Lee (1998). Most free-living larger foraminifers host algal endosymbionts in a relationship analogous to that in zoox anthellate corals (Lee and Anderson 1991). Unlike corals, which have exclusively dinoflagellate symbionts, larger foraminifers host a variety of symbiont taxa including chlorophytes, rhodophytes, diatoms, and dinoflagellates (Lee and Anderson 1991). Although less than 10 % of extant families of the class Foraminifera host algal symbionts, these families account for substantial carbonate produc tion (Lee and Anderson 1991). Globally, benthic symbiontbearing foraminifers account for roughly 0.5% of the total annual carbonate production (Langer 1997). Algal symbioses offer several possible advantages to foraminifers. Host foraminifers may utilize end products of symb iont photosynthesis as an energy source (Muller 1978, Hallock 1981). The chemical changes in the cell matrix caused by photosynthesis may enhance calc ification rates in foramini fers (Duguay and Taylor 1978, ter Kuile 1991). In low nutrient environments, algal symbionts may utilize nutrient wastes produced by the host foraminifer (H allock 1999). The benefits from symbiosis and the variety of endosymbionts hosted, many with different photosynthetic responses, may have enabled different foraminiferal taxa to adapt to environments with a wide range of light availability (Hallock 1999).

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2 Taxa Studied Five benthic species (Fig. 1) were chosen for this study, three that occur in the Caribbean [ Archaias angulatus (Fichel and Moll), Cyclorbiculina compressa dOrbigny and Amphistegina gibbosa dOrbigny] and two that occur in the Indo Pacific [ Amphistegina radiata ( Fichtel and Moll ) and A lessonii dOrbigny]. Archaias angulatus and C. compressa bear chlorophyte symbionts while Amphistegina spp. host diatom symbionts (Lee and Anderson 1991). Fo raminiferal taxa that bear chlorophyte symbionts, which include the Archaiasinae (Family Soritidae), are more diverse in the Caribbean region than in the western Paci fic (Hallock 1999). Among the rotaliid families, including the Amphisteginidae, diversity follows the trend seen in many other organisms; that is, higher dive rsities are observed in the IndoPacific region than either the central Pacific or the western Atlantic/Caribbean (Hallock 1999). Archaias angulatus is the shallowest dwelling of the species studied (<1m ~30m). They are commonly found in abundance in shallow tropical marine environments often in association with Thalassia testudinum Konig sea-grass beds (Duguay 1983). They are sensitive to hypoxia a nd require sufficient water circulation to maintain permanently oxygenated conditions. On the other hand th eir reticulopodia are weak and therefore A. angulatus are most abundant in relatively low energy environments (Hallock and Peebles 1993). Strong positively phototaxic and negatively geotaxic behavior are likely involved in re solving that apparent paradox. Densities of A. angulatus have been observed as high as 15 x 104 individuals m-2 in the Florida Keys, where they can produce approximately 60 g CaCO 3 m-2yr-1 (Hallock and others 1986).

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3 Figure 1a-f Photographs of all species studied. a. Archaias angulatus b. Cyclorbiculina compressa c. Amphistegina lessonii d. Amphistegina gibbosa e. Amphistegina radiata 1mm 1mm 1mm 1mm 1m m a b c d e

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4 eyi of producing more oxygen than the host/symbiont associations consume es y lgal ents because onas oraminifers are found on a variety ing Archaias angulatus hosts the chloro phyte endosymbiont Chlamydomonas hedl Lee, Crocket, Hagen and Stone (Lee and others 1974). Although soritid foraminifers are capable (Kanwisher and Wainwright 1967, Muscatine 1973, Taylor 1973), A. angulatus deriv less than 10% of its organic carbon from the symbionts (Lee and Bock 1976). A majorit of its organic carbon requirements come from grazing and A. angulatus will quickly begin to digest its endosymbionts if no food source is available (Hallock and Peebles 1993). Light has been shown to enhance calcif ication in this speci es (Duguay and Taylor 1978). Cyclorbiculina compressa inhabits slightly deeper waters than A. angulatus typically 5 40 m depth. They are found in hi ghest concentrations in filamentous a mats in open reef environments. They are able to live in higher energy environm they embed themselves in this algal mat (Hallock and Peebles 1993). Lutze and Wefer (1980) found densities of C. compressa to be approximately 200/m2 in Harrington Sound, Bermuda. Cyclorbiculina compressa hosts the chlorophyte symbiont Chlamydom provasoli Lee, McEnery and Kahn (Lee and others 1979). As in A. angulatus, C. compressa will not calcify without its symb ionts or in the dark (Duguay 1983). Amphistegina gibbosa is the smallest and deepest living of the western Atlantic species examined. They can be found from de pths less than 1 m down to 100 m, but are most abundant at depths of 15-40 m (Hallock 1999). These f of reef substrates including coral rubble, phytal substrates and sandy environments (Hallock 1999). Amphistegina gibbosa host diatom symbionts belong

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5 some k osts diatom ke the lent relatively abundant trophic resources (Hallock and Peebles 1993). Therefore, the primary role of chloro phyte symbnts may be enhancement of calcification and secondari 3). In the low nutrient environments inhabited by amphisteg inids, diatom endosymbionts can provide an organic carbon source that is in limited supply (Hallock 1999). The ability of diatoms to several genera, though Nitzschia frustulum var. symbiotica Lee and Reimer emend. appears to be the dominant symbiont in A. gibbosa from the Florida Keys (Lee and others 1995, Lee 1998). Many of the lesser symbionts are rare in natural communities and have been found only as symbionts (Lee and others 1989). Amphistegina lessonii is considered the Pacific equivalent of A. gibbosa (Halloc and others 1996), although A. lessonii have a slightly shallo wer depth distribution (Hallock 1999). These foraminifers are mo st abundant at depths of 10-30 m (Hallock 1984, Hohenegger 1994). In shallo w, high light environments, A. lessonii avoid damage from intense light by cryptic behavior (Hallock 1999). Amphistegina lessonii h symbionts belonging to se veral genera (Lee and others 1993). Amphistegina radiata is a deeper dwelling Pacific amphisteginid (Hohenegger 1994). These foraminifers are found in greate st concentrations from 20-50 m depth and have been found alive as deep as 100 m (Hohenegger 1994). Morphologically they are larger and have a more biconvex shape than either A. gibbosa or A. lessonii Li other two amphisteginids, they host diatom symbionts (Lee and others 1993). In general, soritid foraminifers inhabit shallower, higher light environments than the amphisteginids (Hallock 1999). Paradoxica lly, symbiosis is a more important source of energy for the amphisteginids (ter Kuile and others 1987). The soritids are preva in environments with io ly as a food source (Hallock and Peebles 199

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6 to utiliz ls, photosynthesis exceeds respiration in shallow, well illumin (~ ption of oxygen. Oxygen in liquid environments has been measured by several differen h amic e light in the blue-green range allo ws their hosts to exploit deeper habitats (Leutenegger 1984). Estimating Photosynthesis and Respiration In zooxanthellate cora ated, tropical waters (Muscatine 1990). The same is true for some chlorophytebearing foraminifers (Kanwisher and Wa inwright 1967, Muscatine 1973, Taylor 1973). Lee and others (1980) demonstrated net primary production in Amphistegina lobifera Larsen, a species similar to the foraminifers used in this study, at light levels of 10 klx 180 mol photon m-2sec-1). Photosynthesis or respiration can be gauged by measuring the production or consum t techniques. Early researchers us ed the Winkler method (Winkler 1888), whic has gone through modifications to improve pr ecision (Bryan and others 1976). This method is still in use, though response times are too slow for applications in dyn environments where organisms are respiri ng or photosynthesizing (Gatti and others 2002). With the development of polarographic techniques, measurement of changing oxygen concentration in the liquid phase becam e more reliable. Polarographic study of oxygen led to the development of an oxygen electrode for the study of oxygen concentrations in blood samples (Clark 1956). The resultant probe became known as the Clark-type electrode.

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7 ode de connected by an electr olyte bridge. When a polarizing voltage is applied to the cell e in ith the anode placed in a circul ar well that acts as a reservoir for the electrol a semialso ghtii te tion of the annual carbonate production in shallow tropical mari ne environments (Langer 1997). They The Clark-type oxygen electrode is an el ectrochemical cell that has its cath and ano ionization of the el ectrolyte induces current flow through the electrode. Th magnitude of the current flow is proportional to the concentration of dissolved oxygen the electrolyte soluti on. The concentration of oxygen in the electrolyte is in turn proportional to the oxygen in the surroundi ng environment (Hansatech Instruments 2000). The Hansatech Oxygen Electrode Disk is a Clark-type oxygen electrode developed by Delieu and Walker (1981). The cathode and anode are embedded in an epoxy disk w yte solution. A large cathode is lo cated on top of a small dome where it is covered with a spacer, which provides a uniform layer of electrolyte solution, and permeable membrane. The oxygen electrode provides a stable reading of oxygen concentrations without the labor intensive pr ocedures required by Winkler methods (Gatti et al. 2002). Oxygen electrodes have been extensively used to study metabolic responses of organisms. Lees and others (1991) a nd Catonguy and Markhart (1991) used the Hansatech instrument to quantif y photosynthesis in terrestrial plants. The instrument has been used successfully to mon itor photosynthesis and respiration in Halodule wri Aschers and Thalassia testudinum (Neely 1996 and Berns 2003). Foraminifera are important contributors to benthic communities. They contribu to sediment production (Hallock 1981), accounti ng for a significant frac

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8 occupy f many other photos y ptake n at light intensities as high as 1000 mol photon m-2sec-1. ) mphistegina lessonii were sh res to s in great abundance, tropical marine environments from the shore line to the depths of the euphotic zone (Hohenegger 1994). Despite extensive studies o aspects of the biology and ecology of foramini fers with algal symbionts, there has not been extensive study of their photosy nthetic and respiratory responses. Duguay and Taylor (1978) recorded pr imary production and calcification in Archaias angulatus Using photosynthetic carbon fixation as a measure of ynthesis, they reported that A. angulatus reached light saturati on at a light intensit of ~ 200 mol photon m-2sec-1. Duguay (1983) reported th at calcium and carbon u approach saturation in A. angulatus between 250-500 mol photon m-2sec-1 and showed no photoinhibitio Using differential manometer systems, Lee and others (1980) measured oxygen evolution in Amphistegina lobifera and A. lessonii, among other foraminifers, and observed photoinhibition at light intensity values of 20 klx (~ 360 mol photon m-2sec-1in A. lessonii Using radiocarbon techniques, rates of carbon fixation in A own to differ significantly between fo raminifers incubated in the light and those incubated in darkness (Muller 1978). Others that have used radioc arbon procedu study calcification and productivity include Smith (1977), Spero and Parker (1985), Gastrich and Bartha (1988), and Leutenegger and Hansen (1979). Other studies have examined photosynthesis and respiration rates in foraminifer using micro-sensors to measure O 2 CO 2 pH, and Ca2+ at the test surface of the planktonic foraminifer Orbulina universa, which host dinoflagellate endosymbionts (Rink and others 1998), and the benthic species Amphistegina lobifera and Marginopora

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9 7 to 22.7 d at light intensities as high as 2000 d others (1998) recorded significant increase in O. universa respiration rates masured in the light over va lues measured in the dark. Hannah and others (1994) show -Diver micr orespirometry, that some non-symbiont bearing foraminifers respire es th at of similar sized naked a ves oxygen electrode for the stu foramin a) s oraminifers; mon benthic foraminifera, Archaias angulatus Cyclobiculina compressa Amphistegina gibbosa A. lessonii and A. radiata ; and vertebralis (Rink and Kuhl 2000, 2001). Net photosynthesis rates between 3. nmoles O 2 foraminifer-1 hr-1 were recorded in Amphistegina lobifera Kohler-Rink an Kuhl (2001) also calculated I k values, which approximate light saturation, for A. lobifera of 95 mol photon m-2sec-1 and did not observe photoinhibition mol photon m-2sec-1. Kohler-Rink and othe rs (1998) recorded net photosynthesis rates of 5.3 +/2.7 nmol O 2 h-1 foraminifer-1 in Orbulina universa In addition, Rink an e ed, using Cartesian d at rates ten tim moeba. Thesis Objecti Adaptation of the methods used with the Clark-type dy of plant tissue provides a tool to i nvestigate physiologic responses in larger ifers. The objectives of the research were to: develop techniques using a Hansatech DW1 Oxygen Electrode Unit to asses photosynthesis and respiration in larger f b) estimate photosynthesis and respiration ra tes, photosynthesis/ respiration ratios and generate photosynthesis/irradian ce curves for five com

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10 c) estimate photosynthetic contri bution of these la rge foraminifers to the benthic communities using abundance data frm previous studies. o

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11 red free in plastic jars for transport. Upon arrival at e laboratory the samples were again rinsed with seawater and tran sferred to large petri a 12-hour lighedule at 2 ncubation intensity wa ly 5 mol photon m-2s fter approximaly 24 hoursnisms f the sedimet where they are eily located und icroscope and removed with forceps for use in experimtal procedures rchaias angulatu ens were also collected from Florida Bay waters directly behind the Keys Marine Laborator located in L Long Key (Forg ed ask a of w Methods Collection and Storage of Specimens Four species of large benthic foraminifers, Cyclobiculina compressa Amphistegina gibbosa Amphistegina lessonii and Amphistegina radiata, were collected by diving using SCUBA in the Florida Keys (F ig. 2) and Papua New Guinea at Ambitile Island (Fig. 3). Pieces of coral rubble were collected under water, placed in zippe plastic bags and taken to a fi eld laboratory or shipboard wh ere the rubble was bushed of foraminifers, sediment and other debris. The organic debris and fine sediment was removed as effectively as possible by decanting from the samples. The remaining sediment and foraminifers were then placed th dishes for storage in an environmental chamber. They were kept under ht/dark sc 5 C. I light s approximate ec-1. A te the orga move to the surface o n as e r a stereo m en A s specim y ayton, Florida, on ig. 2). The anisms were collect using m nd snorkel in 1-2 meters ater.

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12 Handful-size clumps of Thalassia testudinum (seagrass) blades and filamentous algae were collected into zippered plastic bags. Because Archaias pseudopodial attachment is not strong, agitating the sample was usually enough to shake th e foraminifers loose from the phytal substrate. The foraminifers and a ssociated sediment were then transferred to plastic bags that were to pped off with oxygen for transport. A sample of the T. testudinum and filamentous algae was also transfe rred to a separate bag for transport. Upon arrival at the laborat ory, the sediment and the Thalassia samples were reunited in a small aquarium and an air stone was added to provide the sample with adequate dissolved oxygen. The aquarium was placed in the lab under ambient light levels (approximately 3-10 mol photon m-2sec-1 depending on time of day and cloud cover. Archaias angulatus are intolerant of low dissolved oxyge n levels (Hallock and Peebles 1993). Transporting the foraminifera separate from the Thalassia sample with oxygen in the ing elow the tolerance level of the organism. Table 1. Dates, locations, and depths of collection of foraminifers used in this study. Specimen Species Date Collection site Depth sample bag prevented dissolved oxygen levels in the foraminiferal sample from dropp b AA01-AA03 Archaias angulatus 5/7/2003 Florida Bay, Keys Marine Laboratory1-2 m AA04 11/1/2003Long Key, Florida Keys AA05-AA10 1/18/2004 CC01-CC10 Cyclorbiculina compressa 12/1/98 Conch Reef, Florida Keys 30 m AG01-1-AG10-5 Amphistegina gibbosa 1/17/2004Tennessee Reef, Florida Keys 10 m AL01-1-AL05-3 Amphistegina lessonii 11/3/03 Tatum Bay, Ambitle Island, 20 m Papua New Guinea Tatum Bay, Ambitle Island, 20 m Papua New Guinea AR01-AR10 Amphistegina radiata 11/3/03

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Figure 2. Collection sites of Archaias angulatus (KML), Cyclorbiculina compressa (Conch Reef) andAmphistegina gibbosa (Tennessee Reef). KML is the Keys Marine Laboratory Figure 3. Location of Ambitle Island, Papua New Guinea 13

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14 leeve mber to ll-mixed water sample. The entire apparatus is placed in a dark container (a he sleeve al algae ere e reaction chamber just above the stir bar. Because Photosynthesis and Respiration Trials Photosynthesis and respiration rates were measured using the Hansatech DW1 Oxygen Electrode Unit. The unit has a Cla rk-type oxygen electrode at the bottom of a 3 ml reaction chamber. A sleeve that allowe d water to be circulated around the chamber to maintain constant temperature surrounded the chamber. The chamber and water s assembly rested on a magnetic stirrer that drove a magnetic stir bar inside the cha maintain a we cardboard box lined with black construction paper). Two holes were cut in the box: one to allow for passage of the water hoses from the temperature-controlled bath to t and a second to provide a w indow for light to enter. The setup procedure included a control r un to determine if the instrument sign exhibited any drift. If a ch ange in the signal was detected during the blank trial, the electrode was cleaned and reassembled prio r to running trials with foraminifers. Individual foraminifera we re picked from the sample dishes under the microscope or hand picked from the aquarium. They were prepared by removing all debris and that were attached to the test or held by rh izopodia. Small paintbrushes and forceps w used to clean the organisms (Duguay a nd Taylor 1978, Lee and others 1980). For the larger species, C. compressa, Archaias angulatus and Amphistegina radiata, individual organisms were placed in a small mesh envelope made from fiberglass screen and suspended in th of the smaller relative size of A. lessonii and A. gibbosa, multiple specimens were

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15A. ter he /I trials run on the first species tested, Cyclorbiculina oped. The water bath that l was also recorded. For the other species, new water bath with no measurable temper ature fluctuations was used and P/I trials ere run for 10 minutes. Respiration rates were estimated with th e foraminifers suspended in the reaction hamber in the dark. As soon as the respira tion trial was complete, photosynthesis trials required to produce changes measurable by the instrument. Three A. lessonii and five gibbosa were used per trial. A cardboard cover was placed over the window and a lid was put on the top of the box to create a dark space. In additi on, black plastic was placed over the box to prevent any light from entering the box th rough cracks around the openings for the wa hoses. For each trial, the specimen (or gr oup of specimens) was left in the dark for approximately one hour to allo w it to stabilize and to a ssure that respiration was measurable. All photosynthesis vs. irradiance (P/I) tria ls were run with the temperature of t water bath set to 25 C. P compressa, were conducted with an older water bath that had a temperature variation of up to +/0.2 C. Since oxygen electrodes are in general as sensitive to temperature change as they are to change in O 2 concentrations, a method to remove the fluctuations in the signal caused by the unstable water ba th was devel regulated the temperature of the reac tion chamber fluctuated between 24.8 C and 25.2 C when set to 25 C. This fluctuation was quite regular and completed 3 cycles in approximately 11 minutes. The voltage readi ngs were recorded when the water bath reached a temperature of exactly 25.0 C as the value was climbing on the third cycle. The exact time elapsed from the start of the tria a w c

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16 began. Photosynthesis trials wer t intensity to the highest in increasing order. The light source employed was the Hansatech LS2 unit, which uses a y data y hesis, 3) light-saturated photosynthesis s ty trial, the light source was turned off and a second respiration trial was run in darkness. After photosynthesis and resp re complete, the organisms were moved from the chamber and measured for maximum, intermediate and minimum diamete e run from th e lowest ligh tungsten quartz halogen bulb with a typical spectrum of approximately 300 750 nm and provides a uniform field of illumination. Light intensity was varied for the P/I trials b altering the distance of the lig ht source and using screens and a set of neutral density filters to attenuate the light. Light intensities were chosen to allow for at least three points to be taken in each of the following range s: 1) dark respiration to approximatel the compensation point, 2) light-limited photos ynt and photo-inhibition. These rang es were determined by running preliminary trials at various light intensities. The varying light intensity trials were run in approximately 11-minute intervals a described above for Cyclorbiculina compressa and 10-minute intervals for all other species. After completion of the highest light intensi iration tria ls we re rs under a microscope. The specime ns were then blotted dry, weighed, and placed on a filter pad, wrapped in al uminum foil and stored at -39 C until chlorophyll extractions were performed.

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17 e n a allowed to warm t s. fficiency Light in g Chlorophyll Extraction Chlorophyll extraction and measurement was performed following the methanol procedure of Holm-Hansen and Rieman ( 1978). Specimens were removed from th freezer and placed whole in cuvettes containing 5 ml of methanol. The cuvettes were covered with para-film to prev ent evaporation of the methanol and then wrapped in aluminum foil to keep them in the dark during extraction. Samples were placed o shaker in a refrigerator and agitated for 18 hours. After 18 hours the samples were removed from the refrigerator and o room temperature. Chlorophyll concentration in the methanol was then measured for each sample using one of two fluorometers. For the Cyclorbiculina trials an older Turner Designs Model 10 unit was used to determine chlor ophyll concentration For all other species a Turner 10 AU unit was used. Data Analysis To generate maximum p hotosynthesis values (P max ) and photosynthetic e ( ), photosynthesis/respiration and irradiance data were evaluated by fitting the oxygen data to the hyperbolic tangent equation described by Jassby and Platt (1976): P = P max tanh ( I/ P max ) tensity (I k ) corresponding to P max was determined by dividing P max by Regressions were run on photosynt hesis/irradiance data for all species normalized to chlorophyll a and mm2 surface area using SPSS Inc. Sigma Plot 5.0 statistical

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18 0. ht t the raw ere nthesis and respiration. The net O2 production or consumption repr esents the combination of symbiont photosynthesis and both symbiont and foraminifer respiration. When software. Surface area is defined as the area in mm2 of the upper surfac e of the organism in question. Descriptive statis tical analyses were performe d using Microsoft Excel 200 To determine if differences existed between mean photosynthesis values at specific lig intensities, two-tailed Student t-tests (Z ar 1984) were performed. At higher ligh intensities, where photosynthesis values appa rently declined significantly, multiple onetailed Students t-tests (Zar 1984) were perfor med starting at the highest light intensity. T-tests were performed on data from successively lower light intensities until no significant difference was found. Photosynthesis values from th ese light intensities were then used in linear regressions to determine photo-inhibition values ( ). Rates of oxygen consumption or production in the dark and at 12 light intensities were measured for each species of foraminife rs to estimate respiration and photosynthesis rates and to construct phot osynthesis/irradiance (PI) curv es. Maximum photosynthesis (P max ), photosynthetic efficiency ( ), and irradiance (I k ) corresponding with maximum photosynthesis values were determined from th e PI curves (Fig. 4) generated by fitting data from individual foraminifers or groups of foraminifers to the hyperbolic tangent equation described a bove (Jassby and Platt 1976). Individual organisms w used in photosynthesis irradiance trials for Archaias angulatus, Cyclorbiculina compressa and Amphistegina radiata. Limitations of the instrument, as well as relative small size and corresponding low oxygen produc tion rates required th e use of groups of three A. lessonii and groups of five A. gibbosa per trial. Oxygen production and consumption was us ed as a measure of photosy

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19 measuring oxygen production or consumption in phytoplankton, the dark respiration is often minimal and is quickly overtaken by oxygen production and positive values are recorded at low light intens ities. When the oxygen consumption of the foraminifer is added to the oxygen consumption of th e symbiont, oxygen production must be considerably higher to overta ke respiration. Because of the added oxygen consumption, net O2 production is not observed until the organism is exposed to higher light intensities and negative values are recorded at lower light intensities. However, a decrease in oxygen consumption is observed at the lower light intensities. To fit the data sets to the Jassby and Platt (1976) equation, prior to r unning regressions on the raw data sets, pretrial dark respiration rate s were removed by subtracting th e respective dark respiration rate from all data points in th e sets to generate gross photosyn thesis rates. Raw values of oxygen production represent net photosynthesis. Photosynthesis/Irradiance curves and the corresponding values for Pmax, and Ik were determined using gross photosynthesis rates. ly. Oxygen consumption rates were recorded be fore light intensity trials began and immediately after the highest light intensity trial. Metabolic scope, the difference between pre and post trial resp iration rates, was calculated for all species. Metabolic scope is an indicator of the organisms ability to increase metabolic activity in pursuit of food or other survival strategies. In addition, metabolic factorial scope, the ratio of post trial to pre-trial oxygen production rate was ca lculated. Metabolic factorial scope is often used in place of metabolic scope when comparing different organisms. Figure 4 shows a sample P/I curve, with P max , and I k illustrated graphical

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20 Figure 4. Sample photosynthesis/irradiance curve showing P max , I k and parameters. SpecimenIrradiance Phynthesis -200100120140160 Pmax 02004006008001000 Ik 20406080 otos

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21 RESULTS Archaias angulatus Rates of oxygen consumption and production were measured in ten Archaias angulatus specimens (Fig. 1a) collected from the Florida Keys (Fig. 2, Table 1). The physical parameters of the individual foramini fers are summarized in Table 2. All were relatively large specimens, ranging from 2.15 to 4.25 mm in maximum diameter, with upper surface areas estimated at 3.25 mm2 10.5 mm2. Masses ranged from 1.06 to 3.85 mg. The amounts of chlorophyll a extracted from single in dividuals were 0.190 g 1.18 g. The quantity of chlorophyll extracted from Archaias angulatus appears to be highly correlated, however specimen AA08 was la rger than all other Archaias specimens. If this specimen is removed from the data analysis, chlorophyll a concentration is not correlated to either mass or surface area (Fig. 5a,b). Table 2. Physical parameters of Archaias angulatus Parameter Range Median MeanStandard Deviation Major Diameter (mm) 2.15-4.25 2.90 2.87 0.56 Intermediate Diameter (mm) 1.9-3.15 2.23 2.24 0.37 Minor Diameter (mm) 0.35-0.55 0.53 0.47 0.10 Upper Surface Area (mm2) 3.20-10.5 5.11 5.19 2.05 Mass (mg) 1.06-3.85 1.74 1.86 0.79 Chlorophyll a Extracted (g) 0.190-1.180.46 0.49 0.29

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22 Mass Vs. Chl a Archaias 0.0000.2000.6000.800Mass of Foram (mg)Ch 0.4001.0001.2001.4000246l a extracted(ug) Surface Area Vs. Chl a Archaias 0.0000.2000.600Surface Area (mm2)Ch(ug) 0.4000.8001.0001.2001.400051015l a extracted a. b. angulatus Oxygen consumption rates were measured prior to the start of ately one hour (Appendix A; Table A-3, A-4). intensity of 0.96 t oxygen production at the lowest light level (Appendix A; Table A-3, A-4). net production of oxygen was observed at all higher light intensities in all individuals (Fig. 6, 7). Maximum hotosynthetic rate (Pmax) ranged from 54.8 to 202 nmoles O2 hr-1g chl a-1 (5.57 to 14.3 moles O2 hr-1 mm-2) (Table 3, 4). Archaias angulatus reached Pmax at irradiances nging from 35.2 to 163 mol photon m-2sec-1 (Table 3, 4). Eight of the ten specimens xhibited highest O2 production at 542 mol photon m-2sec-1 and nine of the ten showed Figure 5. Correlation of mass (a) and surface area (b) to Chl a extracted from individual Archaias photosynthesis/irradiance trials and after the foraminifers were acclimated to the reaction chamber in the dark for approxim Seven of the A. angulatus showed net oxygen consumption at the lowest light mol photon m-2sec-1. The oxygen consumption rate at the lowest light intensity exceeded the rate recorded in dark trials for five of the individuals. The other three individuals showed ne After the initial increase in oxygen consumption by half the specimens, oxygen production rapidly increased as light intensity increased, and p n ra e

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23 x A; Table A3). Post-illumination respiration measu owat la four-focrease in en consumption over the initia l dark trialls, wit etabolic factorial a-1 n respiration was 25..51 nles r-1 g Ch nmoles O2 hr-1 mm-2). Mean post illumination oxygen consumption was 121 +/18.5 2 2ption ally within 30-4 nation levels when the protists eft in thear Photoinhibition was observed at li ensitean 542 hoton m-ues r m toth a mean of -0.043 +/.033 when normalized to Chl a and 0.009 +/.003 when normalized to surface area (Table 3, 4). the completion of light trials. The center of the foraminife rs had a normal green coloration while the perimeter of the organisms had no coloration. marked decrease in oxygen production at higher light intensities (Appendi rements sh ed east ld in oxyg leve h a mean m scope of 9.6 +/5.7 moles O 2 hr ug chl -1 (Table 3, 4). Mean oxygen consum ption l evels in pre-illuminatio 0 +/6 mo O 2 h l a-1 (1.43 +/0 n moles O hr-1 g Chl a-1 (7.25 +/0.552 nmoles O hr-1 mm-2). This higher consum rate declined asympt otic 0 min to pre-illum i were l dk. g ht int ies gr ter tha mol p 2sec-1 in most individuals. Beta val anged fro 0.100 0 wi 0 Specimens of Archaias showed redistribution of sy mbionts at

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24Table 3. Derived photosynthetic parameters normalized to Chl a for Archaias angulatus Parameter Range Median Mean Standard Deviation Pmax (nmoles O2 mg chl a-1) 54.8-202 120.5 122 46.2 Alpha 0.55-5.74 1.2 1.7 1.6 Ik (umol photon m-2sec-1) 35.2-163 85.7 95.6 38.6 Beta -0.1 0 0 0 Initial Respiration (nmoles O2 mg chl a-1) 0-43.1 9.2 13.6 12.7 Post Trial Respiration (nmoles O2 mg chl a-1) 50.2-172 88.6 95.5 35.5 Metabolic Factorial Scope 4 21 9 9.6 5.7 Table 4. Derived photosynthetic para meters normalized to surface area for Archaias angulatus Parameter Range Median Mean Standard Deviation Pmax (nmoles O2 mm-2) 5.57-14.3 9.84 10.20 2.69 Alpha 0.074-0.28 0.11 0.12 0.06 Ik (umol photon m-2sec-1) 35.2-163 85.70 95.10 38.50 Beta -0.01 0.00 0.00 0.00 Initial Respiration (nmoles O2 mm-2) 0-2.05 0.94 0.99 0.65 2.08 etabolic Factorial Scope 4 21 9.00 9.60 5.70 Post Trial Respiration (nmoles O 2 mm-2) 5.30-12.3 7.42 8.00 M

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-50 200 25 0 502 100 1500200400600800100012001400Light Intensiy (umole photon m-2 s-1) g Chl a -1 nmoles O hr-1 Figure 6. Photosynthesis vs irradiance normalized to Chl a for Archaias angulatus

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-4 26 -2n 0240200400600800100012001400Light Intensiy (umole photon m-2 s-1)ms O2 hr g Chl a-1 681012-1 ole Figure 7. Photosynthesis vs irradiance normalized to surface area for Archaias angulatus Cyclorbiculina compressa Rates of oxygen consumption and production were measured in ten Cyclorbiculina compressa specimens collected from the Florida Keys (Fig. 1, 2, Table 1). The physical parameters of the individual foraminifers are summarized in Table 5. All were relatively large specimens, ranging from 3.5 to 5.6 mm in maximum diameter, with 1 7.7 mg. The amounts of chlorophyll extracted from single individuals were 0.181 g 0.799 upper surface areas estimated at 9.21 mm2 24.2 mm2. Mass wet weight ranged from 2. g The quantity of chlorophyll extracted from Cyclobiculina compressa was

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27of highly correlated to surface area and mass of the individual organism with r2 values 0.99 and 0.97 respectively (Fig. 8 a, b). Table 5. Physical parameters of Cyclorbiculina compressa Parameter Range Median MeanStandard Deviation Major Diameter (mm) 3.5-5.6 3.9 4.24 0.77 Intermediate Diameter (mm) 3.25-5.5 3.63 4.03 0.81 Minor Diameter (mm) N.A. N.A. N.A. N.A. Upper Surface Area (mm) 9.19-24.211.1 13Mass (mg) 2.-7.7 3 3. 2.8 5.5 177 1.95 Chlorophyll a Extracted (g) 0.18-0.800.241 0.3880.241 Mass Vs. Chl a Cyclo rbiculina0.2000.600 0.000C 0.400(ug) 0.8001.000 05Mass of Foram (mg) 10 hl a extracted S ual a 0.0.00.0.10ce Area rface Are Vs. Ch Cyclorbiculina 000 200 .400 600800 .000 0 102Surfa 03(mm2) C hl a extracted (ug) igure 8. Correlation of mass (a) and surface area (b) to Chl a extracted for C. compressa Oxygetion rates were ed trt riance trails and afti wclimated to the reaction r approximately o Aihowed itial ither mghyll or mmace areaendix B: Eight individuals exhibited net consumption of oxygen when exposed to the lowest light intensity of 0.96 mol photon m-2 sec-1. The respiration rate at the lowest a. b F n consump measur prior to he sta of photos ynthesis/irad er the foram nifers ere ac chamber in the dark fo ne hour. ll spec mens s low in respiration rates normalized to e chlorop 2 surf (App Table B-3, B-4).

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28e not light intensity exceeded the rate recorded in dark trials for six of the individuals. No changes in oxygen concentrations were observed for one specimen, while one specimen exhibited net oxygen production at this light intensity. Although the mean net respiration rate for this light intensity was slightly higher than the initial dark rate, the rates wersignificantly different (Fig. 9,10). -4 0 -20020nmoles O2-1 4060800510152025303540Light Intensiy (umole photon m-2 s-1) hr g Chl a-1 Archaias Cyclobiculina igure 9. Mean oxygen production/consumption normalized to Chl a showing initial increase in oxygen consumption at lowest light intensity in Cyclorbiculina compressa. F

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-2-10123450510152025303540Light Intensity (umole photon m-2 s-1)nmoles O2 hr-1 g Chl a-1 Archaias Cyclobiculina Figure 10. Mean Oxygen production/consumption normalized to surface area showing initial increase in After the initial increase iny most specimens, oxygen production rapidly increased as light intensity increased, and net production of oxygen was oblina f oxygen consumption at lowest light intensity in Cyclorbiculina compressa. oxygen consumption b served at all higher light intensities in all individuals (Fig. 11, 12). Derived parameters are summarized in tables 6 and 7. Maximum photosynthesis (P max ) ranged from 115 189 nmoles O 2 hr-1g chl a-1 (2.39-5.03 nmoles O 2 hr-1 mm-2). Cyclorbicucompressa reached P max at irradiances between 70.8-160 mol photon m-2sec-1. Seven othe ten specimens exhibited highest O 2 production at 542 mol photon m-2sec-1, and all specimens showed a marked decrease in oxygen production at light intensities higher than 779 mol photon m-2sec-1 (Appendix B). 29

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30 Range Median Mean Standard Deviation Table 6. Derived photosynthetic parameters normalized to Chl a for Cyclorbiculina compressa Parameter Pmax (nmoles O2 mg chl a ) 115-189 140.20 144.00 22.30 -1 Alpha .918-1.831.23 1.27 0.30 I (umol photon m-2sec-1) 70.8-160 122.00 119.00 30.50 .23 -0.07 -0.08 0.07 itial Resoles O2 mg c-11 17.60 nmoles O27-157.00 69.70 -68.70 18.50 k Beta -0 In piration (nm hl a ) 0-5 4.8 5.20 19.90 Post Trial Respiration ( mg chl a-1) 10 316 181.00 Metabolic Factorial Scope 4.2 2.5 15.80 Table 7. Der ived photosynthe tic ord tce area fo lorbiculina compressa Parameter Range Median Mean Standard Deviation p ara meters n malize o surfa r Cylc P max (nmoles O2 mm-2) 2.39-5.22 3.84 3.73 1.00 A lpha .0176-.05670.03 0.03 0.01 Ik (umol photon m-2sec-1) 70.8-160 122.00 119.0030.50 Beta 0.00 0.00 0.00 0.00 Initial Respiration (nmoles O2 mm-2) 0-1.08 0.37 0.48 0.40 Post Trial Respiration (nmoles O2 mm-2) 2.71-6.06 4.45 4.40 0.93 Metabolic Factorial Scope 4.2 -62.5 8.70 15.80 18.50 Post-illumination measurements showed al most an order of magnitude increase in ial scope was 5.8 +/18.5. Mean oxygen consumption levels in pre-illumination respiration were 19.8 +/5.6 Mean postin 30-40 -1. oxygen consumption over the initia l dark trial levels. Mean me tabolic factor 1 nmoles O 2 hr-1 g Chl a-1 (0.48 +/0.125 nmoles O 2 hr-1 mm-2). illumination oxygen consumption was 181 +/22.0 nmoles O 2 hr-1 g Chl a-1 (4.4 +/2.9 nmoles O 2 hr-1 mm-2). The higher consumption rate dec lined asymptotically with min to pre-illumination levels when the protists were left in the dark. Photoinhibition was observed at light intensities above 779 mol photon m-2sec

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31t er of the individual, while the outer most chambers of the test were colorless, resultin All individuals studied showed a redistribution of symbionts during the experimental trials, which lasted for 126 min. At the onset of the photosynthesis/irradiance trials, the foraminifers were a uniform green color throughoutheir tests. Upon completion of the trials, the green color was concentrated around the cent g in a white ring around the perimeter of the protist. -40-200 20nmoles O 4060040068121400Light Intensitye pm-2 hr 80100120140160-1 g Chl a-1 020 00 00 1000 00 (umol hoton 2 s-1) s irradiance normalized tr Ccuompressa. Figure 11. Photosynthesis v o Chl a fo yclorbi lina c

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-1-101 5 1 22 2 hr-1 3 g Chl a 34001400Light Intensity (umole photon m-2 s-1)nmoles O 4-1 0200400600800100012 Figure 12. Photosynthesis vs irradiance normalized to surface area for Cyclorbiculina compressa. Amphistegina gibbosa Rates of oxygen consumption and production were measured in ten groups of five individual Amphistegina gibbosa specimens collected near Tennessee Reef, in the Florida Keys (Table 1, Figure 2). The physical parameters of the individual foraminifers are summarized in Table 8. Size ranged from 0.75 to 1.4 mm in maximum diameter, with upper surface areas estimated at 0.38 mm2 to 1.37 mm2. Masses ranged from 0.10 to 0.83 g. The amounts of chlorophyll extracted from single individuals were 0.025 to 0.143 g. The quantity of chlorophyll extracted from Amphistegina gibbosa was highly m n 32

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33correla and ted to surface area and mass of the individual organism with r2 values of 0.880.83 respectively (Fig. 13 a, b). Table 8. Physical parameters of Amphistegina gibbosa Parameter Range Median MeanStandard Deviation Major Diameter (mm) 0.75-1.40 1.1.04 0.17 03 Intermediate Diameter (mm) 0.65-1.25 0.93 0.92 0.14 Minor Diameter (mm) 0.25-0.65 0.40 0.42 0.08 Upper Surface Area (mm2 ) 0.38-1.37 0.76 0.76 0.23 Mass (mg) 0.10-0.83 0.36 0.34 0.16 Chlorop hyll a Extracted (g) 0.025-0.1430.06 0.07 0.03 Mass Vs. Chl a A. gibbosa0.0000. 05000.20.40.60.81Mass of Foram (mg) 0.000.1500.200Chl a extracted (ug) 1 Surface Area Vs. Chl a A. gibbosa0.000 0.200 0.05000.511.5Surface Area (mm2)Chl 0.1000.150 a extracted(ug) ion l ion at the lowest light intensity of 0.96 mol photon m-2sec-1. Oxygen consumption rate exceeded the rate corded in dark trials for four of these groups. Three groups showed no change in net a. b. Figure 13. Correlation of mass (a) and surface area (b) to Chl a extracted for A. gibbossa. Oxygen consumption rates were measured prior to the start of photosynthesis/irradiance trails and after the foraminifers were acclimated to the reactchamber in the dark for approximately one hour. All groups showed low initiarespiration rates normalized to either mg chlorophyll or mm2 surface area (Appendix C). All of the groups of A. gibbosa showed net oxygen consumpt re

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34 oxygen p at the lowt l ex net oxygen grd be the cosation point where serv of ghnsities ( ndix C: Table C-3, CDerived parameters for A. gibbosa are summarized in tables 9 and 10. Maximum photosynthesis (Pmax) ranged from 14.8 to 58.5 nmoles O2 hr-1g chl a-1(1 nmoles O2 hr-1 mm-2). Amphistegina gibbosa reached Pmax at irradiances from 9 -27 mol photon m-2 sec-1. Photosynthesis dramatically incr eased in all groups between 3 and 17 mol photon m-2sec-1, even in those that showed net consumption of oxygen throughout their trial. At light intensities between 17 and 779 mol photon m-2sec-1, oxygen production Table C3, C4). able 9. Derived photosynthetic parameters normalized to Chl a for Amphistegina gibbosa Parameter Range Median Mean Standard Deviation roduction est ligh eve l and three hibite d decreased consumption. Five of the oups faile to clim abov mpen oxygen production is ob ed at any the li t inte Appe 4). either plateaued or fluctuated in all groups (Appendix C T P max (nmoles O 2 mg chl a-1) 14.8-58.634.50 36.00 14.90 Alpha 1.48-2.091.85 1.80 0.24 chl a-1) 43.8-104 65.00 68.50 19.30 1.1 14 1.90 3.77 4.04 I k (mol photon m-2sec-1) 8.95-27.213.20 14.30 5.44 Beta -0.05 -0.05 -0.05 0.01 Initial Respiration (nmoles O 2 mg chl a-1) 5.48-69.627.50 33.90 24.40 Post Trial Respiration (nmoles O 2 mg Metabolic Factorial Scope

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35 Range Median Mean Standard Deviation Table 10. Derived photosynthetic parameters normalized to surface area for Amphistegina gibbosa Parameter Pmax (nmoles O2 mm) 0.971-4.13 3.12 3.19 1.34 -2 Alpha I (mol photon m-2sec-1) 8.96-27.2 13.20 14.30 5.45 -2 0.0996-0.3710.23 0.23 0.09 k Beta 0.00 0.00 0.00 0.00 Initial Respiration (nmoles O 2 mm) 0.440-6.55 2.50 2.99 2.17 Post Trial Respiration (nmoles O 2 mm-2) 4.34-7.64 5.85 5.87 1.15 Metabolic Factorial Scope 1.1 14 1.90 3.80 4.00 -50 -40-30030400400Light Intenlen)nmoles O2 evolved hr-1 -20-10102050 ug Chl a-1 A. lessonnii A. gibb 200 600 800 1000-2 -1 12001400 siy (umo photo ms osa A. radiata 14. Photosynthesis vs irradiance normalized tofor tpecies Figure Chl a he amphisteginid s

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-6-4-20 36 2-1 46ns -2 mm A. le ssonnii A. g ibbosa A. r evolved hr O2 0200400600800100012001400Light Intensity (umole photon m-2 s-1)mole adiata on respiration rates were almost double pre-trial rates. Mean metaboles O2 oxygen consumption was 68.5 +/6.1 nmoles O2 hr-1 g Chl -1 (5.9 +/0.4 nmoles O2 hr-1 mm-2) (Table 9, 10). This higher consumption rate declined asymptotically within 30tion levels when the protists ere left in the dark. -1. Figure 15. Photosynthesis Vs Irradiance normalized to Suface Area for the amphisteginid species Post illuminati lic factorial scope was 3.8 +/4.0. Mean oxygen consumption levels in pre-illumination respiration was 33.9 +/7.7 nmoles O 2 hr-1 g Chl a-1 (3.0 +/0.7 nmohr-1 mm-2). Mean post illumination a -40 min to pre-illumina w Photoinhibition was observed at light intensities above 542 mol photon m-2secAt the conclusion of the experimental trials most individual foraminifers appeared slightly pale and uneven in color.

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37 inifers Individual s ranged from 1.0 to 1.5 mm in maximum diameter, with upper surface areas estimated at 0.71 mm2 to 1.53 mm2. Masses ranged rom 0.26 to 1.03 mg. The amounts of chlor ophyll extracted from single individuals hyll extracted from individual nii was h e a idual f 0.0.93ec (Fig. 16. o a i Parameter Range Median MeanStandard Deviation Amphistegina lessonii Rates of oxygen consumption and production were measured in five groups of three individual Amphistegina lessonii specimens collected near Ambitile Island in Papua New Guinea (Table 1, Figure 3). The physical pa rameters of the individual foram are summarized in Table 11 f w ere 0.076 0.228 g. The quantity of chlorop Amphistegina lesso ighly correlated to s urfac rea ass of the indiv nd ma organism with r va 2lues o 94 and resp tively a, b) T able 11. Physical parameters f Amphistegin lessoni M ajor Diameter (mm) 1.0-1.5 1.20 1.22 0.02 Intermediate Diameter (mm) 0.90-1.35 1.05 1.07 0.14 Minor Diameter (mm) 0.45-0.70 0.55 0.58 0.08 Upper Surface Area (mm2) 0.71-1.53 0.98 1.05 0.27 Mass (mg) 0.22-1.03 0.51 0.53 0.26 Chlorophyll a Extracted (g) 0.072-0.2280.12 0.13 0.05

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Mass Vs. Chl a A. lessonii0.10.20.25 (ug) 38 0Mass of Foram (mg) 0.050.1500.511.5Chl a extracted Surface Area Vs. Chl a A. lessonii0.2000.250 0.000Surface Area (mm2)Ch 0.0500.1000.15000.511.52l a extracted(ug) a. b. ated to the reaction chamben at the lowest light inest light Figure 16. Correlation of mass (a) and surface area (b) to Chl a extracted for A. lessonii Oxygen consumption rates were measured prior to the start of photosynthesis/irradiance trails and after the foraminifers were acclim r in the dark for approximately one hour. All specimens showed low initial respiration rates when normalized to either mg chlorophyll or mm2 surface area (Appendix D, Table D3, D4). All of the groups of A. lessonii showed net oxygen consumptio tensity of 0.96 mol photon m-2sec-1. Oxygen consumption rate at the lowintensity exceeded the rate recorded in dark trials for two groups. The other three groups showed no change in net oxygen production at the lowest light level. The mean net O 2 flux for this light intensity was slightly less than the initial dark rate but the rates are not significantly different. After the initial increase in oxygen consumption by most groups of specimensoxygen production rapidly increased as light intensity increased (Fig. 14, 15), and net production of oxygen was observed at all higher light intensities up to 779 mol photon m-2sec-1. At 1,288 mol photon m-2sec-1, four of the five groups fall back into net oxygen consumption. Derived parameters for A. lessonii are summarized in tables 12 and 13.

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39 2 -2). Amphistegina lessonii reached Pmax at irradiances from 21.2 to 29.6 mol photon m-2sec-1, and 22.2 to 32.2 mol photon m-2sec-1 when ormalized to surface area. All groups rapidly increased oxygen production to 36 mol hoton m-2sec-1 then fluctuated above that intensity, and exhibited maximum oxygen roduction at 542 mol photon m-2sec-1 followed by marked decrease in oxygen hetic parameters norml a le eter M Standaiation Maximum photosynthesis (P max ) ranged from 39.0 to 61.8 nmoles O 2 hr-1g chl a-1( 5.0 to 6.48 nmoles O 2 hr-1 mm n p p production there after (Fig. 14, 15). Table 12. Derived photosynt a lized to Ch for A. ssonii Param Range edian Mean rd Dev P max ( nmoles O2 mg chl a-1) 384 9.0-61. 1.80 46.00 9.20 Alpha 1.50-4.26 2.37 2.57 0.94 I k (umol photon m-2sec-1) 2 itial Respiration (nmoles O2 mg chl a-1) 4.10-32.318.30 19.40 11.50 ost Trial Respiration (nmoles O2 mg chl a-1) 44.8-104 73.90 74.80 23.00 etabolic Factorial Scope 2.8 18 3.40 6.20 6.60 1.8-29.6 26.40 25.50 3.33 Beta -0.04 0.05 -0.05 0.01 In P M synthetic para meters no surea A. lessonii r MMStandarion Table 13. Derived photo rmalized to face ar Paramete Range edian ean d Deviat P max ( nmoles O2 mm-2) 5.0 55 0. 2-6.48 .79 .73 68 Alpha 0 00 0. 2222 4. -0-0 0. itial Respiration (nmoles O2 mm-2) .569-4.11 2.18 2.31 1.30 Post Trial Respiration (nmoles O2 mm-2) 6.29-11.3 10.20 9.09 2.12 etabolic Factorial Scope 2.8 18 3.40 6.20 6.60 .185-0.256 .23 .22 03 I k (umol photon m-2sec-1) .2-32.2 7.10 6.40 12 Beta 0.00 .01 .01 00 In M

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40 al .582 his higher consumption rate dec ved at light intensities above 542 mol photon m-2sec-1. At the conclusion of the experimental trials most individual foraminifers appeared slightly pale and uneven in color. Amphistegina radiata Rates of oxygen consumption and producti on were measured in ten individual Amphistegina radiata specimens collected near Ambitile Island in Papua New Guinea (Table 1 Figure 1, 3). The physical parame ters of the individual foraminifers are summarized in Table 14. Size ranged from 1.70 to 2.15 mm in maximum diameter, with upper surface areas estimated at 2.14 to 3.38 mm2. Masses ranged from 0.98 2.12 mg. The amount of chlorophyll extracted from single individuals was 0.196 0.590 g. The quantity of chlorophyll extracted from Amphistegina radiata was highly correlated to surface area and mass of the in dividual organism with r2 values of 0.87 and 0.85 respectively (Fig. 17 a, b). Post illumination respiration rates were, on average, 3-fold higher than pre-tri rates. Mean metabolic fact orial scope was 6.2 +/6.6. M ean oxygen consumption levels in pre-illumination respiration was 19.4 +/5.16 nmoles O 2 hr-1 g Chl a-1 (2.31 +/0 nmoles O 2 hr-1 mm-2). Mean post illumination oxygen consumption was 74.7 +/10.3 nmoles O 2 hr-1 g Chl a-1 (9.09 +/0.949 nmoles O 2 hr-1 mm-2). T lined asymptotically within 30-40 min to pre-illumination levels when the protists were left in the dark. Photoinhibition was obser

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41 Table 14. Physical parameters of Amphistegina radiat Parameter Range Standard Deviation a Median Mean Major Diameter (mm) 1.7-2.15 0.13 1.80 1.84 Intermediate Diameter (mm) 1.6-2.0 1.70 1.72 0.12 Minor Diameter (mm) 0.6-0.8 0.68 0.69 0.06 Upper S2Mass (m urface Area (mm) 2.13-3.38 2.44 2.50 0.35 g) 0.98-2.12 1.54 1.57 0.29 Chlorophyll a Extracted (g) 0.196-0.5900.30 0.32 0.11 Mass Vs. Chl a A.radiata00.20.60123Mass of Foram (mg)l 0.40.8Ch a extracted (ug) Surface Area Vs. Chl a A. radiata0.0000.2000.60001234Surface Area (mm)l 0.4000.8002Ch a extracted (ug) a. b. Figure 17. Correlation of mass (a) and surface area (b) to Chl a extracted for A. radiata photosynthesis/irradiance trails and after the foraminifers were acclimated to the reaction Oxygen consumption rates were measured prior to the start of chamber in the dark for approximately one hour. All specimens showed low initial respiration rates normalized to either g chlorophyll or mm2 surface area (Appendix E, Eight of the A. radiata showed net oxygen consumption at the lowest light -2-1ndividuals. One individual showed no change in net oxygen production at the lowest light level and three exhibited decreased net oxygen Table E-3 and E-4). intensity of 0.96 mol photon msec. Oxygen consumption rate exceeded the rate recorded in dark trials for six of these i

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42 consum ld be fit A. o ax at irradiances from 7.5 to179 lized to um increase in the rate of photosynthesis, represented by maxim ion 1 ption. Six of the groups failed to climb above the compensation point, where oxygen production is observed, at any of the light intensities tested. When normalized to g Chl a only seven of the 10 data sets produced cou to the Jassby and Platt (1976) equation. Wh en normalized to surface area only six could be fit. Values for oxygen consumption and production were much more variable forradiata then either of the other amphi steginids or the soritid species. Derived parameters for A. radiata are summarized in Tables 15 and 16. Maximum photosynthesis (P max ) ranged from 4.31 49.2 nmoles O 2 hr-1g chl a-1(1.68 t 3.98 nmoles O 2 hr-1 mm-2). Amphistegina radiata reached P m mol photon m-2sec-1, and from 7.3-190 mol photon m-2sec-1 when norma surface area. Despite the variability, eight sp ecimens showed an increase in rates of net oxygen production or a decrease in ne t oxygen consumption between 0 and 11 mol photon m-2sec-1. The maxim um increase in oxygen production or maximum decrease in oxygen consumpt is evident in the first four light intensities ranging from 0.96 .9 mol photon m-2sec-1All specimens fluctuated in net oxy gen production or consumption to 779 mol photon m-2sec-1, then oxygen consumption increased at 1,288 mol photon m-2sec-1 in 9 of 0 specimens (Fig. 14, 15).

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43 eter Range MedianMean Standard Deviation Table 15. Derived photosynthetic parameters normalized to Chl a for Amphistegina radiata Param Pmax (nmoles O2 mg chl a-1) 4.31-49.2 17.10 21.40 15.50 Alpha 0.0698-3.161.27 1.29 1.25 Ik (umol photon m-2sec-1) 62.30 Beta 0.02 Initial Respiration (nmoles O2 mg chl a-1) 9.31-79 30.30 31.40 17.80 Post Trial Respiration (nmoles O2 mg chl a-1) 0-132 73.40 69.60 46.00 Metabol 7.49-179 13.50 48.90 -0.06 -0.04 -0.04 0. ic Factorial Scope 0 4.7 2.30 2.40 1.40 on Table 16. Derived photosynthetic parame ters normalized to surface area for Amphistegina radiata Parameter Range Median Mean Standard Deviati Pmax (nmoles O2 mm-2) 1.68-7.10 2.65 3.31 2.04 Alpha 0.0275-0.3650.21 0.19 0.14 I (umol photon m-2sec-1) 7.26-191 12.20 48.70 72.50 -2 k Beta -0.01 0.00 0.00 0.00 Initial Respiration (nmoles O 2 mm ) 1.33-9.92 3.53 3.94 2.45 Post Trial Respiration (nmoles O 2 mm-2) 0-18.5 9.22 8.99 4.86 Metabolic Factorial Scope 0 4.7 2.30 2.40 1.40 Post illumination respiration rates were almost double pre-trial rates. Mean metabolic factorial scope was 2.4 +/1.4. Mean oxygen consumption rates in preillumination respiration were 31.4 +/5.64 nmoles O hr-1 g Chl a-1 (3.94 +/0.7 nmoles O hr-1 mm-2) (Table 3,4). Mean post illumination oxygen 276 2 consumption was 69.6 +/14.52 2E, nmoles O hr-1 g Chl a-1 (8.99 +/1.54 nmoles O hr-1 mm-2) (Appendix Table E-3, E-4). This higher consumption rate declined asymptotically within 30-40 min to pre-illumination levels when the protists were left in the dark.

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44-1. s most individual foraminifers appeared slightly Photoinhibition was observed at light intensities above 542 mol photon m-2sec At the conclusion of the experimental trial pale a nd uneven in color.

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45 nts are bearing taxa are present fr om shallow, high light environments (Hallock and Peebles 1993) to the light-lim ited depths of the photic zone (Hohenegger 1994). Their success as carbonate producers is notable in the geologic record at times llock 1987) and organic production uch as two orders of ma gnitude (Bralower and Theirstein, 1984). Variou tic electrodes makes them useful tools for mea field Discussion Benthic foraminifers thrive in many di fferent ecological ni ches (Murray 1991). They are found living in sediment, on shells and rocks, and on plants (Murray 1991). Their distribution in shallow tropical waters includes environments where nutrie limiting and environments where trophic reso urces are abundant (Hallock and Peebles 1993). Symbiontwhen oceanic circulation was diminished (Lee and Ha was reduced as m s metabolic parameters, including respiration, endosymbiont photosynthe capacity and response, and metabolic scope pr ovide some clues to the strategies of metabolic adaptation used by foraminifers to exploit a wide range of habitats. Photosynthesis and respiration can ef fectively be estimated using oxygen electrodes. The quick response times of C lark-type suring oxygen production in water. They are adaptable to a variety of environmental chambers and are relatively inexpensive. The Hansatech DW1 Liquid Phase Oxygen Electrode Unit is easily transpor table and can be easily set up in most laboratories where power is available.

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46 sent all changes in oxygen concentration in relatively small volumes of water. The large platinum cathode on the Hansatech electrode is no exception. As the electrode consumes oxygen, the concentration immediately above the electrode declines. The sea-water must be adequately stirred to compensate for the consumption of oxygen at the surf ace of the electrode. If the water in the chamber is not adequately stirred, the instrument signal will slowly drift dow n. The signal will also drift if the speed of the stir-bar is too great. Th is results in a time consuming, trial and error procedure for identifying the opt imum speed for the stir bar. Selection of a water bath for use with the instrument is also an important consideration. As observed in early experiments, oxygen electrodes are very sensitive to temperature changes. As a consequence a wa ter bath with temperature fluctuation less than 0.1 C should be used. Phytoplankton have the ability to ph otoacclimate when cultured under light intensities outside their optim um range (Richardson and others 1983). Therefore, light intensity at which the foraminifers were main tained may have influenced responses. An argument might be made that, by incubati ng the foraminifers at 5 mole photons m-2 s-1, the organisms used in the experiments were preconditioned to low light intensities prior to experimental trials. The procedur e (e.g., Talge and Hallock 2003, Williams and Hallock 2004) of maintaining Amphistegina species at intensi ties < 10 mole photon m-2 s-1 emerged from recurring observations in the early 1980s that exposure to higher intensities induced bleaching (H allock and others 1986). Since Cyclorbiculina compressa individuals were picked from sedime nt samples incubated under low light Clark-type oxygen electrodes consume oxygen at the cathode. This can pre problems when measuring sm

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47 ic kton typically exhibit net oxygen production at very low light levels. Incorporati ng the oxygen consumption by the foraminifers with the oxygen prod symbiont resulted in negative values for oxygen production at the lowest light intensity tested, 0.96 mole photon et oxygen s tensity. Dark oxygen consumption accounts for consum ow in conditions, it is possible that these speci mens were also preconditioned prior to experimental trials. Photosynthesis in Symbiotic Associations P max is defined as the maximum rate of photosynthesis attained by the organism It was calculated by fitting gross photosynthesi s and irradiance values to the hyperbol tangent equation [P = P max tanh ( I/ P max )] described by Jassby and Platt (1976). Photosynthesis/Irradiance curves generate d for phytoplan uction and consumption of the m-2 s-1, and in the trials using Amphistegina gibbossa and A. radiata, n production was seldom attained. Therefore gross photosynthesis values were used to calculate photosynthetic parameters including P max I k and Gross photosynthesis rate were calculated by removing dark oxygen consumption rate from the net oxygen production rates at each light in ption by both the foraminifer and the endosymbionts. Light requirements for growth and p hotosynthesis have been shown to be significantly different between different al gal classes of phytoplankton (Richardson and others 1983). Diatoms can survive and grow at very low photon flux densities, yet can tolerate relatively high light intensities. In contrast, chlorophytes are unable to gr

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48dson and others 1983). very low light environments and reach max photosynthesis rates in much higher light intensities (Richar The soritid foraminifers, Archaias angulatus and Cyclorbiculina compressa, and the amphisteginid foraminifers, Amphistegina lessonii, A. gibbosa and A. radiata, show similar photosynthetic/irradiance responses to the free-living taxa represented by their symbionts. When normalized to chlorophyll a, the chlorophyte-bearing soritids reachhigher P max values than do the diatom-bearing amphisteginids (Fig. 18 20). -50050100150 200 ug Chl a-1 0200400600800100012001400Light Intensiy (umole photon m-2 s-1)nmoles O2 evolved hr-1 Archaias Cyclobiculina A. lessonnii A. radiata A. gibbosa Figure 18. Photosynthesis vs irradiance normalized to Chl a for all species

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49 -6Light Intensity (umole photon m-2 s-1) -4-2024100200400600800100012001400nmoles O2 evolved hr-1 Archaias Cyclobiculina A. lessonnii A. gibbosa 68 mm-2 A. radiata ecies examined. Intermediate values were observed in the um photosynthetic rates may have several potential causes including differences in photosynthetic response of the diatom symbionts and differences in the metabolic rates of the foraminifers. Figure 19. Photosynthesis vs irradiance normalized to surface area for all species Among the amphisteginids there is a gradation of oxygen production rates, with the highest rates observed in the A. lessonii, the shallowest-dwelling sp A. gibbosa, which Hallock and others (1986) found to be less light tolerant than A. lessonii and which exhibits a slightly deeper distribution (Hallock 1999). The lowest P max values were observed in A. radiata (Fig. 20), the foraminifer with the deepest depth distribution. Differences in their maxim

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020406080100120140160A.angulatusC. compressaA. lessoniiA. gibbosaA.radiataForaminiferal SpeciesO2 Evolved (nmol hr-1 ug chl a-1) Figure 20. Maximum photosynthesis (P) normalized to Chl a The three amphisteginid species likely posses different suites of diatom endosymbionts, with the deeper dwelling species possibly utilizing symbionts that are adapted to lower light intensities (Lee and others 1980). Generally, low light adapted phytoplankton species exhibit lower P values than those seen in species adapted to higher light intensities (Richardson and others 1983). When normalized to surface area, the P values for Cyclorbiculina compressa are similar to those observed in Amphistegina spp. (Fig. 21). This is most likely due to a lower concentration of chlorophyll a per unit mass in C. compressa. Chlorophyll a concentrations measured in C. compressa are only 40 % of those measured in A. angulatus and A. lessonii and only half the concentrations seen in A. gibbosa and A. radiata (Fig. 22). max max max 50

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51 0.0002.0006.0008.000angulatompressA. lessA. gibbosA.radiaForaminiferal Species2 Evolved hr 4.00012.000A.usC. caoniiataO-1 mm 10.000-2 Figure 21. Maximum photosynthesis (P max ) normalized to surface area 0 0.050.10.150.20.250.3AraiasCyclorulinaA. soniiA. bosaA.adiataForaminiferal Species chbiclesgib rChl. a (ug Chl.a mg specimen) Figure 22. Mean chlorophyll a concentrations for each species. imens era Symbiont distribution appears uneven in Cyclorbiculina compressa. Spectypically exhibit a scalloped pattern (Fig. 1b), where some areas of the foraminif

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52 d ikely x values fonii ). d fitting the trend observed where shallow sis increases with respect er appear bright green and other areas appear pale. Although individual C. compressa ten to be larger than the A. angulatus, they have signifi cantly lower chlorophyll a concentrations (Fig 22). Differences in chlorophyll a concentration per unit mass are also the most l explanation for the differences observed in P max within the amphisteginids. When normalized to surface area, P max values for A. lessonii are significantly higher than P ma or either A. gibbosa or A. radiata (Fig. 21). Chlorophyll a concentrations observed in the amphisteginids correspond to the observed P max values with A. lesshaving a significantly higher concentration than either A. gibbosa or A. radiata (Fig. 22 P max values for the amphisteginid species match up very well with the values recorded by Kohler-Rin k and Kuhl (2001) for A. lobifera when their values are converte and normalized to surface area. Calculat ed gross photosynthesis value, expressed by oxygen production, for A. lobifera was 8.2 +/1.6 nmoles O 2 hr-1 mm-2. This rate is higher than the rate calculated for A. lessonii (Fig. 21) er dwelling species showing higher P max values. Photosynthetic Efficiency ( ) The value is the slope of the light-lim ited portion of the P/I curve. Photosynthetic efficiency represents the ra te at which photosynthe to irradiance (Falkow ski 1997). Although chlorophytes generally reach high maximum photosynthetic rates, they typically do not reach P max as quickly as diatoms (Richardson and others 1983).

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53 Calculated values for specim atically higher than all other A. angulatus species and were higher than most values calc is value is removed, A. angulatus values are also significantly different than Amphistegina lessonii and A. cies. Therefore it is difficult to make conclusions based on this data set, and median d n for all three amphisteginid sp ecies (Fig. 24). When including all data points, d ig. For data normalized to chl a, calculated values for C. compressa are significantly lower than values for Amphistegina lessonii and A. gibbosa (Fig. 23). Alpha values for Archaias angulatus include an outlier in the data set. en AA03 were dram ulated for the am phisteginid species. If th gibbosa. Not all data sets for A. radiata adequately fit the Jassby a nd Platt (1976) curve. In addition, the variability in alpha va lues is considerably higher for A. radiata than for the other spe values may be more meaningful than means. Although raw data values an derived parameters values are variable in A. radiata, the species show a decrease in oxygen consumption between 0 and 17 mol photon m-2sec-1. For data normalized to surface area, values for C. compressa are significantly different tha alpha values for A. angulatus are not significantly different than the amphistegini values. However, when the outlier is removed, the values are significa ntly different (F 24).

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3.03.5 0.00.51.01.5.5rchaiarchaiasCyclo. less.gibbo. radiSpeciesAlpha 2.02As IA IIrAoniiAsaAata Figure 23. Alpha values for all species normalized to Chl a. Archaias I data includes all 10 samples. Archaias II is the same data set excluding specimen AA03, which was an outlier. 0.000.050.100.200.2530rchaiasrchaiasCycl. less.gibbo. radiSpecies 0.150.A IA IIorAoniiAsaAataAlpha Figure 24. Alpha values for all species normalized to surface area. Archaias I data includes all 10 samples. Archaias II is the same data set excluding specimen AA03, which was an outlier. 54

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55 al the values observed in the amphis al al milar whethe-1 Irradiance Measures I k is a calculated value based on and represents the intersection of the initi light-limited portion of the P/I curve and P max (Fig. 4). The soritid species reached P max at significantly higher irradiance values (I k ) than teginids (Fig. 25). This response agai n reflects the general ch aracter of the alg endosymbiont taxa, as chlorophyte algae gene rally exhibit higher photosynthetic potenti than that observed in diatoms (Richardson a nd others 1983). These values remain si r the original data are normalized to Chl a or surface area. Photosynthesis/irradiance curves can change due to photoacclimation (KohlerRink and Kuhl 2001). The overall higher I k values observed by those researchers may be due to different pre-trial maintenance conditions, which were ~5 mole photon m-2sec for the three amphisteginids and C. compressa and ~10 mol photon m-2sec-1 for A. angulatus. These values are probably lowe r than in-situ light intensities.

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0 140) 4060 20A.eC. comsaA. loniiA. aA. radiataSpecies 80100120 angulatupresessgibbosIk (umole photons m-2 s-1 Figure 25. Irradiance values (I k ) at maximum photosynthesis for all species As is observed in P max values, I k values are significantly different between A. lessonii and A. gibbosa (Fig. 25), with lower values recorded for A. gibbosa. Hothers (1986b) reported that growth rates in A. lessonii and A. gibbosa are similar when allock and grown t -2-1t than A. gibbosa, they apparently prefer environments with significantly different light intensity. If the two species were to at lower light intensities but at 40 mole photon m-2sec-1, A. lessonii had a higher growth rate. Growth rate in A. gibbosa reaches saturation at light intensities of 6-8 mole photon m-2sec-1 and the organisms show increased frequency of bleaching at higher lightintensities (William and Hallock, 2004). Talge and Hallock (2003) recorded significandeterioration of symbionts and endoplasm in A. gibbosa at light levels as low as 13-15 mol photon msec. Previous studies have demonstrated that A. gibbosa has a low lightolerance; the results from my study support those observations. Although A. lessonii only exhibit a slightly shallower depth distribution 56

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57 exist in the same geographic area, there ma significant difference in their depth ranges. r locations (Lee and others 1980). Since th e foraminifers were collected from dissonii ole photon m-2sec-1 (Fig. 25). Duguay 50 ss 542 -1, which is more consistent with values reported by Duguay (1983). Based o ted y be a more Amphistegina gibbosa is known to utilize endosym bionts that are adapted to relatively low light intensities (Lee and others 1980), allowing the pr otists to exploit low light environments. Although individual amphi steginid specimens collected from the same environment possess very similar suites of algal endosymbionts, there can be variability in the complement of the endosymbionts when specimens are collected from different depths o fferent environments, it is possi ble that the differences seen between A. leand A. gibbosa are due to the different photosynthe tic responses of the unique suite of symbionts or the different metabolic responses of the foraminifers. Using oxygen production as a measure of photosynthesis, values of I k for Archaias angulatus were calculated to be 96 +/12 m (1983) reported maximum calcium and carbon uptake in A. angulatus at 200-2 mole photon m-2sec-1. I k values are inherently lower than the intensity at which maximum photosynthesis is observed. I k values are calculated from P max and when paired with fitting the data to the hyperbolic tangent equation, results in values for I k le than raw data values observed. Raw data indicated maximum oxygen production at mole photon m-2sec n the I k values calculated, Archaias angulatus in shallow water is light satura for most of the day (Duguay 1983). Duguay and Taylor (1978) found carbon fixation to be light limited up to intensities of ~200 mole photon m-2sec-1 in A. angulatus and that there was no

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58 6 mole photon m-2sec-1. Lee attributed this difference in photosynthetic behavior to the adaptation of diatom pigm ent to lower light intensities and shorter wavelengths than that of zoochlorellae (Lee a nd others 1980). Observed differences of Ik ohler-Rink and Kuhl (2001) reported onset of light saturation at levels of 164198 m ids uld be ce area, t. Additional trials would have to be run with this species to reduce the error be s significant difference in carbon fi xation at light levels of 380 mole photon m-2sec-1. Hallock and others (1986) observed photoinhibition in Amphistegina lessonii at light levels below between soritids and amphisteginids in this study agree with these findings (Fig. 25). K ole photon m-2sec-1 for the dinoflagellate bearing foraminifer Amphisorus hemprichii and 95 mole photon m-2sec-1 for the diatom bearing species Amphistegina lobifera. Although the values they calculated are significantly higher than the values recorded in this study, I k values are higher for the forami nifers bearing symbionts from higher light adapted taxa. The differences of response observed by Kohler-Rink and Kuhl (2001) in I k values is similar to that seen be tween the chlorophytes -bearing sorit and the diatom-bearing amphi steginids in this study. Variability of I k values was substantially higher in A. radiata than in all other species studied. When normalized to chlorophyll a, only 7 of the 10 data sets co successfully fit to the Jassby and Platt (1976) equation. When normalized to surfa only 6 of the 10 sets successfully fit the curve. For this reason the values observed for A. radiata are suspec fore any hard conclu sions could be reached on I k values. Calculated mean and median values for I k are very similar for Amphistegina gibbosa and A. lessonii. Mean I k value for A. radiata of 49 mole photon m-2sec-1, i dramatically higher than values seen in the other amphisteginids. However, median I k

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59 values may be a better indicator of the I cteristic s of the raw photosynt hesis to irradiance data that are masked when the data sets ar e fit to the hyberbolic tangent equation. These changes represent sig several of the species values for A. radiata (12-13 mole photon m-2sec-1) are very similar to those for A. gibbossa (13 mole photon m-2sec-1). Thus the median k parameter, in this case of high individual variability. Photosynthesis and Irradiance Raw Data There are several distinct chara nificant excursions in the data sets and are observed in Both the foraminifers and their al gal symbionts consume oxygen through metabolic processes. In the current study there is no way to separate the metabolic contribution of the host from that of the alga l symbiont. In additi on, respiration rates for all the species change during the course of the trials from low initial levels to high post trial levels. In 5 of 10 Archaias angulatus specimens and 6 of 10 Cyclorbiculina compressa specimens, oxygen consumption was greater at the initial light intensity (0.96 mole photon m-2sec-1) than at the initial da rk trial (Appendix A, Table A-3; Appendix B, Table B-3). The initial increase in oxygen consumption by C. compressa is illustrated in Figures 9 and 10. Although the mean oxygen c onsumption values are not significantly different, there appears to be a metabolic change in either the symbiont or the foraminifer as the symbiosis ramps up photosynthesis. While 7 of 10 A. angulatus individuals

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60 es cant in Amp as ity observed in the data for this species. in s in mbers. y. decrease in oxygen produ ction at higher light levels may be due to increased metabolic activity by the foraminifer as it attempts to relocate to a shaded show an increase or no change in oxygen consumption the mean value reflects net production (Fig. 9, 10). All three amphisteginid species exhibited similar fluctuations in the P/I curv (Fig. 14, 15). Oxygen production peaked at three intensities 36.5, 175 and 542 mole photon m-2sec-1, with a subsequent declin e in production at the next higher light intensity. The first drop in oxygen production betw een light intensities of 36 and 48 mole photon m-2sec-1 was not statistically significant in any of the three species. The decline in oxygen production recorded between 175 and 233 mole photon m-2sec-1 was signifi histegina lessonii and A. gibbosa when data were normalized to either chlorophylla or surface area. The decline in ox ygen production at thes e intensities in A. radiata w not significant, reflecting the high variabil There are several potential causes of these fluctuations includi ng: 1) increase metabolic rate of the symbiont; 2) increase in metabolic rate of the foraminifers; 3) decrease in oxygen output due to photoinhibition; 4) the pres ence of multiple species of endosymbionts with different light requirements, and 5) multiple layers of symbiont inner chambers of the test. Outer chambers and their resident symbionts may initially shade symbionts in inner chambers of the foraminifers. Light levels in the inner chambers may be significantly less than what is available to symbionts in outer cha As light levels increase and more light penetr ates through the test to the inner chambers, the inner chamber symbionts may increas e their oxygen production accordingl Amphisteginids are known to exhibit cryp tic behavior and are phototaxic (Zmiri and others 1974). The

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61 location for and tion in oxygen produc to ri butions. If these differen t the ng drop ight intensitds what is optimal for survival. No such ability has been described. It is of the cytoplasing from en f ox er nse to changing pholux de nsities. This is beyond the scope of this To facilitate this movement the foraminifers have developed mechanisms the rapid assembly and disassembly of micr otubules, which allow for rapid extension retraction of psuedopodia (Welnhofer and Travis 1996). The reduc ed by the organism could be the result of the increased oxygen consumption due metabolic activity associated with movement. During exposure to higher light intensities, A. gibbosa has been observed to relocate cytoplasmic material into pore c ups (Talge and Hallock 2003). The metabolic cost of this internal mobilization of cytoplasm may also reduce net O 2 production. The three amphisteginid species have di fferent depth dist ces are due to preferences for different light intensitie s, it could be argued tha species attempt to relocate to locations with optimum light availability, including seeki shelter when light intensity exceeds their rela tive optimal light intensities. This may account for one of the three dr ops in oxygen production seen at higher light intensities in the P/I curves generated from raw data. The similarities in intensities at which the in oxygen production occurs for each individual amphisteginid species may be due to the relatively large differences between the hi gher light intensities chosen for the study. Phototaxic behavior and the foraminifers ability to mobilize cytoplasm in response to changing light intensities implies the organisms ability to sense when l y excee possible that the foraminifers are responding to changes in chemical composition m result dosymbionts re lease o ygen and oth photosynthetic products in respo ton f

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62 bilitief th e instrumentation used, howev er it does suggest possible dy Photoinhibition a a process that is timedependant and occurs on the same time scale as is needed to produce P/I curves (Geider and Osborn 1992). The simi onset of photoinhibition may be due to the similar experi mental treatment of the different species. ng species 2001). xposed to nd A. phisorus wer intensities. It is also s study and the capa s o avenues for future stu Photoinhibition was observed in all five species of foraminifera. Archaias angulatus and the three amphisteginid spp. exhibited significant reduction of oxygen production at light intensity levels above 663 mole photon m-2sec-1. Cyclorbiculina compressa exhibited reduction in oxygen production at light intensity levels above 779 mole photon m-2sec-1. Oxygen production data were not recorded for Cyclorbiculin compressa at 663 mole photon m-2sec-1. Photoinhibition is larities in the light intensity values at the Amphistegina lobifera and Amphisorus hempricii, both shallow-dwelli showed no photoinhibition to 2,000 mole photon m-2sec-1 (Kohler-Rink and Kuhl Lee and others (1980) did not measure photoinhibition in these species until e intensities of 3,300 mole photon m-2sec-1. Amphistegina lessonii, A. gibbosa aradiata are all deeper-dwelling species than either Amphistegina lobifera or Am hemprichii and this may account for the apparent phot oinhibition at lo possible that the symbionts photoacc limated to the lower maintenance intensitie and therefore were more readily photoinhibited.

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63 f the uction by the symbionts, following the hyberbolic tangent equation (Jassby and Pla te. post-trial oxygen consumption rate s are significantly different in all five species. Post-illumination rates are as much as a order of magnitude higher than preillumination metabolic rates in the soritids (Fig. 26). Dark respiration rates for all amphisteginid species were comparable to values recorded by Kohler-Rin k and Kuhl (2001) for A. lobifera (3.3 +/0.6 nmoles O2 hr-1 mm-2) and Amphisorus hemprichii ( 0.8 nmoles O2 hr-1 mm-2). Rink and others (1998) recorded both dark respiration rates of 1.7 +/0.7 nmol O2 foraminifer-1 hr-1 and light respiration rates of 3.9 +/nmol O2 foraminifer-1 hr-1 in the planktonic foraminifer Orbulina universa. The metabolic scope is defined as th e difference between the minimum and maximum metabolic rates and is a measure of the total energy an organism can make available for activity or external work (Gordon 1977). Metabolic scope has been used in energetics studies on fish (Cla ireaux and others 2000, Cutts and others 2002, Mallekh and Respiration Respiration rate, represented by oxygen consumption, varied as much as 15 fold from relatively low pre-trials levels to high post irradiance leve ls. The respiratory response at the different light intensities between pre-trial an d post-trial levels cannot be parsed out in this study. It is possible that the foraminifer response shadows that o oxygen prod tt 1976). The pre-trial respiration rate represen ts the foraminifers resting or basal metabolic rate. The post trial rate corresponds to the protists maxi mum metabolic ra Pre-trial and n

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64Lagardere 2002) and other organisms (Staples a others 2002) as an indicator of ability of an organism to increase metabolic activity in the pursuit of food or in other survival strategies. nd 0510152025A. angu latusess C.Species compraA. lessoniiA. gibbosaA. radiataRatio Final Resp/Initial Resp Figure 26. Factors studied Metabolic scope differs significantly between representatives of the two families based on data normalized to chlorophyll a. Cyclorbiculina compressa and Archaias angulatus show higher metabolic scope than the three amphisteginid species. This may be due to a fundamental difference in the feeding strategies between the two taxa. Active organisms tend to exhibit higher metabolic scopes than sedentary ones (Gordon 1977). Archaias angulatus are highly reliant on grazing to provide organic carbon (Lee and Bock 1976, Duguay and Taylor 1978) and both soritid species will digest their endosymbionts when starved (Hallock and Peebles). Conversely the amphisteginids can ial scope (ratio of post-trial respiration rate to pre-trial respiration rate) for all specie

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65 survive in vy endosymbionts darkness for up ly survive a few weeks without feeding. When oxygen data are normalized to surface area, metabolic scope estimates are not distinguishable by family. When comp aring organisms with different metabolic scopes, it is often more effective to comp are their factorial scope, the ratio of active metabolic rate to resting me tabolic rate (Fig. 26) (Gordon 1977). Examining the data in this way shows the soritids with greater metabolic capability than A. gibbosa or A. radiata with A. lessonii values falling in the middle. Rink and others (1998) calcu lated respiration rates of the planktonic species, Orbulina universa to be 3.9 +/1.9 nmole O2 h-1 in the light. Corresponding dark respiration rates were significan tly lower at 1.7 +/0.7 nmole O2 h-1. The calculated metabolic factorial scope of 2.3, is sim ilar to the mean values calculated for A. gibbosa and A. radiata, as well as A. lessonii. Symbiont-bearing Foraminifers as Primary Producers Assuming light intensities of 1,000 mole photon m-2 s-1 at 0.5 m below the surface (Duguay 1983), it is apparent that in shallow water Archaias angulatus spends much of the day in light intensities that are well above saturation levels. Assuming the foraminifers are light saturated for 10 hours a day, an individual foraminifer 1 mm2 in surface area would produce approximately 28 moles O2 yr-1. If there are 104 er low nutrient environments on organic carbon provided by their Amphistegina spp. readily go into a dormant state and survive in to a year (Talge 2002). Archaias angulatus can on

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66 individuals/m2 (Hallock and Peebles 1993), annual yearly O2 production would be more than 280 mmoles O2 m-2 of habitat (Table 17). Even at depths of 18 m where light levels have been recorded at 94 mole photon lge 2002), A. angulatus is very near saturation intensities. Similar calculations onCyclorbiculina compressa indicate annual surface area of f 200 individuals m (Hallock and Peebles 1993). However, ompressa specimens selected for experimental studies were much larger than the average Among the amphisteginids only Amphistegina lessonii showed significant primary oint where oxygen production exceeds c onsumption (Fig. 14, 15). Mean Pmax values for A. radired y Talge (2002) at Conch Reef and densities of 10 individuals m annual primary produc (Table 17). om 28 68 nmoles O2 mg chl a h under saturating light intensities of 330 mole photon Langer and others (1997) suggested that larger foraminifers represent a CO2 source ated CO2 uptake at tion rates th Amphistegina lobifera, indicating that this ecies represents a CO2 sink. Oxygen production rates for A. lessonii exceeded dark s m-2 s-1 (Ta O 2 production to be 9.3 mmoles O 2 m-2 of habitat (Table 17), based on an average 5 mm2 and density o-2 these extrapolations are undoubtably high because individual Archaias angulatus and C c size found in natural populations. production. Amphistegina gibbosa reached P max just above the compensation p ata failed to even reach the compensation poi nt. Using light intensities measu4 -2 b tion for A. lessonii is estimated at 15.3 mmoles O 2 m-2 of habitat Photosynthetic rates in the coral Seriatopera hystrix were determined as ranging fr -1-1 s m-2 s-1 (Burris and others 1983). in reef communities. Rink-Kohl er and Kuhl (2000) demonstr foraminifer shell surface under light conditions and significantly lo wer dark respira an O 2 production rates in light in sp

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67 spiration oxygen consumption at light in tensities greater than 92 mole photons m-2 s-1. Talge ( lobifera (<10 m optimum), (Hallock 1984, Hohenegger 1994). Therefore, Amphiseper, lower light environments, and a s Oxygen production rates for A. gibbosa and A. radiata do not exceed dark respira ource for CO2 on the reef as Langer and others (1997) proposed. spective dark respiration rates at 17.5 mole photons m-2 s-1 and 36 mole photons m-2 s Ma lso significantly xygen roduction and the shallo wer depth distribution, A. angulatus and C. compressa are probab ost of the day and are CO2 sinks. alculated to be appr oximately 3.5 nmoles O2 mm-2 hr-1. These values are similar but slightly ded net O2 roduction per mm gives rates of 5.0 +/1.1 nmoles O2 mm hr-1. Amphistegina lobifern in the data er re 2002) recorded light levels of 94 mole photons m-2 s-1 at depths of 18 m. Amphistegina lessonii have a slightly deeper depth distribution (10-30 m optimum) than A tegina lessonii may be a CO 2 source in de ink in shallow water. tion rates at any of the experimental light intensities, indi cating that they are a s Oxygen production rates in Archaias angulatus and C. compressa exceed their re -1 ximum oxygen production rates for both soritid species are a higher than their pre-trial (dark) oxygen cons umption rates. Because of the high o p ly light saturated for m Net oxygen production rates normalized to surface area for A. lessonii where c lower than values recorded by Kohler-Rink and K uhl (2001) who recor production rates per individual foraminifer in A. lobifera. Converting their values to O 2 p 2 -2 a is the shallowest-dwelling amphisteginid and continues the trend see I recorded for amphisteginid species where by the shallower dwelli ng species show high

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68Amphisand Kuhl 2001). Table 17 stegina lessonii ion oxygen production rates. The va lues are also similar to production rates recorded for orus hemprichii of 3.3 nmoles O 2 mm-2 hr-1 (Kohler-Rink Annual primary production of A. angulatus, C. compressa and Amphi Species Individual O 2 production Field Density Annual O 2 product (moles O 2 individual-1yr-1) (Ind. m-2) (mm oles O2 m yr-1) -2 A rchaias angulatus 28.5 10000 285 (1) Cyclorbiculina compressa 46.5 200 (1)9.3 (2) Amphist 0 15.3 Other Habitats egina lessonii 1.53 1000 Thallasia testudinum 3.02 x 105 (3) 5 (4) Coral re 2.79 x 10 rustose coralline algae 5.4 48 x 103 (5) ef C (1) Hallock and Peebles (1993) (2) Hallo ck 1984 y and others (2002), Onuf (1996) rs (1979) (3) Kald (4) Roge ) Chisholm (2003) Although recorded photosynthesis values for Amphistegina gibbosa hover around the com t intensities test ed when night-time oxygen e used in experimental trials. Use of hi gher resolution instruments would allow more precise eans for measuring oxygen oncentration in solution since Clark (1956) designed the model upon which most (5 pensation point through most of the ligh respiration is considered, this species is a net consumer. The very small changes in production recorded in A. gibbosa stretch the capabilities of the oxygen electrod quantification of the meta bolic needs of this species. Recommendations for Future Research Oxygen electrodes have been a re liable m c

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69 polarographic electrodes is that they consume oxygen at the membrane surface oncentrations, this is no longer a problem (Gatti and others 2002). y enable resolution of unanswe red questions on production in A. gibbosa and A. radiata. produc 8 and 9) and in the fluctuation in oxygen Multiple factors probably contribute to metabolic changes in Amphistegina spp., includi obilization to move to suitable light envi ronments. Further study on what is happening to the i hanges could prove miri 1974), organise responding to and how they a fford this metabolic cost may also prove electrodes are based today. Unfortunately one of the inherent problems with (Hansatech 2000). With the advent of new optical sensors to measure oxygen c Running the photosynthesis/irradiance tria ls using this newer technology ma Higher resolution measurements may also reveal details of the ini tial increase in oxygen tion observed in C. compressa (Fig. production observed in the amphisteginid specie s at higher light in tensities (Fig. 14, 15). ng internal mobilization of cytoplasm (Talge and Hallock 2003) and external m nternal microstructure of the cytoplasm during these c interesting. Since the foraminifers exhibit both positive and negative phototaxis (Z they apparently posses the ability to sen se light. The determination of what the ms ar intriguing.

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70 Maximum oxygen production (P ) when normalized to Chl a was diatom-bearing species. ring A. gibbosa and na compressa, consistent with the ge neral characteristics of Calculated irradiance at P (I ) was estimated at only 13 mole photon indicating very low light requi rements of these species. I value calculated with previous laboratory studies. a compressa were 95 and 119 al hoton m-2 s-1, and in Cyclorbiculina compressa above 779 mole photon m-2s-1. Conclusions max approximately 3-4 fold higher in ch lorophyte-bearing species than in Photosynthetic efficiency ( ) was higher in diatom-bea A. lessonii than in chlorophyte-bearing Archaias angulatus and Cyclorbiculi their symbionts. max k m-2 s-1 in A. gibbosa and A. radiata, consistent with previous studies k for A. lessonii was slightly higher, 26 mole photon m-2 s-1, also consistent I k for Archaias angulatus and Cyclorbiculin mole photon m-2 s-1 respectively, indicating high er light requirements; however these values are lower than previous estimates for optim irradiance. Photoinhibition was observed in Archaias angulatus and all amphisteginids above 663 mole p

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71 rophyte-bearing soritid The ratio of post-trial metabolic rate to pre-trial metabolic rate (metabolic factorial scope) was significantly higher in the chlo species than in the diatom-bearing amphisteginids.

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72 References Berns, D.M., 2003, Physiological responses of Thalassia testudinum and Ruppia maritima to experimental salinity levels: Thesis (M.S.), University of South Florida, 2003, 71p. Brawlower, T.J. and Theirstein, H.R., 1984, Low productivity and the slow deep water circulation in the mid-cretaceous oceans: Geology, v. 12, p. 614-618. Bryan, J.R., Riley, J.P. and Williams, P.J.L., 1976, A winkler procedure for making precise measurements of oxygen concentr ations for productivity and related studies: Journal of Experimental Ma rine Biology and Ecology, v. 21 p. 191-197. Castonguay, Y. and Markhart, A.H., 1991, Saturated rates of photosynthesis in water-stressed leaves of common bean and tepary bean: Crop Science, v. 31 (6), p. 1605-1611. Claireaux, G., Webber, D.M., Largardere, J. P. and Kerr, S.R., 2000, Influence of water temperature and oxygenation on the aerobi c metabolic scope of Atlantic cod (Gadus morhua): Journal of Sea Research, v. 44 (3-4), p. 257-265. Clark L.C., 1956, Monitor and control of blood and tissue oxygen tensions: Transactions American Society for Artificia l Internal Organs, v. 2, p. 41-46. Chisolm, J.R.M., 2003, Primary productivity of reef building crustose coralline algae: Limnology and Oceanography, v. 48 (4), p. 1376-1387. Cutts, C.J., Metcalfe, N.B. and Taylor, A.C., 2002, Fish may fight rather than feed in a novel environment: metabolic rate and feeding motivation in juvenile Atlantic salmon: Journal of Fish Biology, v. 61 (6), p. 1540-1548. Delieu, T.D., and Walker, D.A., 1981, Polargraphic measurement of photosynthesic O2evolution by leaf discs: New Phytologist, v. 89, p. 165-175. Duguay, L.E., 1983, Comparative laboratory and field studies on calcification and carbon fixation in foraminiferal-algal a ssociations: Journal of Foraminiferal Research, v. 13 (4), p. 252-261.

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73 Duguay, L.E., and Taylor D.L., 1978, Primary production and calcification by the soritid foraminifer Archaias angulatus: Journal of Protoz oology, v. 25(3), p. 356-361 Falkowski, P.G., 1980, Light-shade adaptation in marine phytoplankton: In Falkowski, P.G. [Ed.] Primary Productivity in the Sea. Plenum Press, New York, p. 99-119. Falkowski, P.G., 1981, Light-shade adaptati on in marine phytoplankton: Journal of Plankton Research, v. 3 (2), p. 203-216. Falkowski, P.G., Laroche, J., 1991, Acclimation to spectral irradiance in algae: Journal of Phycology, v. 27, p. 8-14. Gatti, S., Brey, T., Muller, W.E.G., Heilm ayer, O. and Holst, G., 2002, Oxygen micro optodes: a new tool for oxygen measurements in aquatic animal ecology: Marine Biology, v. 140, p. 1075-1085. Gastrich, M.D. and Bartha, R., 1988, Primary productivity in the planktonic foraminifera, Globirgerinoides ruber (d Orbigny): Journal of Foarminiferal Research, v. 18 (2), p. 137-142. Geider, R.J., Osborne, B.A., (1992) Algal P hotosynthesis, The Measurement of Algal Gas Exchange. Chapman and Hall, London, England. 256 p Gordon, M.S., 1977, Animal Physiology: Pr inciples and Adaptations: Macmillan Publishing Co., Inc, New York, 699 p. Gupta, B. K., 1999, Modern Foraminifera, Kluwer Academic Publishers, The Netherlands, 371 p. Hallock, P. (1981a) Algal symbiosis: a math ematical analysis. Marine Biology, v. 62, p. 249-55 Hallock, P. (1981b) Production of carbonate sediments by selected foraminifera on two Pacific coral reefs: Journal of Sedimentary Petrology, v. 51, p. 467-74 Hallock, P. (1984) Distribution of larger foraminiferal assemblages on two Pacific coral Reefs: Journal of Foramiferal Research, v. 14, p. 250-61 Hallock, P. (1999) Symbiont Bearing Fora minifera, in Modern Foraminifera (ed. B.K. Gupta) Kluwer Academic Publishers, The Netherlands, 371 p. Hallock, P. and Peebles, M.W., (1993) Fora minifera with chlorophyte endosymbionts: Habitats of six species in the Florid a Keys: Marine Micr opaleontology, v. 20, p. 277-292.

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Hallock, P., Cottey, T.L., Forward, L.B., a 74 sediment production of Arcaias angutus (Foraminiferida) in Largo Sound, Florida: Journal of Foraminiferal Research, v. 16, p. 1-8. es of common benthic foraminifera (protozoa) : Journal of the Marine Biological s n e e ents in the liquid phase. Systems mananh Instr Limorfolk,and 33 p. eggar99istri bof liviner foram of Sesoko-Jima, Okinawa, Japan: Mari nelogy, v.. 291-33 ann1978) Cphyll a mination: improvements in Methodology: Oikos, v. 30, p. 438-447. atical relationship between photosynthesis and light for phytoplankt nology and Oceanography, v. 21 ), p-5 idge, P.M., Cifuentes, L.A. (2003) Carbon budget for a subtropical seagrass dominated coastal la goon: How important are seagrasses to Estuaries, v. 25 (4A) p. 528-539 r, a 67la l in 90 r-Rinku., (2000)osensor studies of photosysnthesis and respiration in laytic foram The po-chemiccroenvironment of Marportebral and Amphisorus hemprichii: Marine Bo v. 137 ( 473-486 photosynthesis and respiration in the ls minifera and Amphisorus hempi,elia, v. 5, p. 111-122 r, M. 9volutionnvi ronmental and econom foramra hrift, Geowissenschaftlichultat Tubingen, 40 raminifera and their endosymbiotic algae: Symbiosis, v. 25 (1-3), p. 71-100 nd Halas, J. (1986), Population biology and la Hannah, F., Rogerson, A., Laybournparry J. (199 4) Respiration rates and biovolum As ociatio n of the U ited Kingdom v. 74 (2), p. 301-312 Hansatech Instrum nts (2000). Oxygen m as urem ual, H satec uments ited, N Engl Hohen J. (1 4). D ution g larg i nifera NW Eco 15, p 4 Holm-Hansen, O. and Riem B. ( hloro deter Jassby, A.D. and Platt, T. (1976) Mathm formulation of the on: Lim (4 540 47. Kaldy, J.E., Onuf, C.P., Eldr total ecosystem net primary producti on? K anwishe Bio J.W., ogical nd Wainwr Bullet ight, SA., (19 v. 133, p. 378-3 ) Oxygen ba nce in some coral reefs: Kohle S, K rger s hl M mbio Micr inifera. I hysic al mi rgino ra ve is, Amphistegina lobifera i logy, 3), p. Kohler-Rink S, Kuhl M., (2001) Microsensor studies of arger ymbiont bearing fora Amphistegrina lobifera richi Oph 5 (2) Lange R., (1 97) E ary, e ical significance of inife Habilitatio ns-sc e Fak pp. Lee, J.J., (1998) Living Sands Larger fo

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75 Lee, J.J and Anderson, O.R. (1991) Symb iosis in foraminifera, in Biology of Foraminifera, (eds J.J. Lee and O.R. Anderson), Academic Press, London, pp157220. Lee, J.J. and Bock, W.D. (1976) The importa nce of feeding in two species of soritid foraminifera with algal symbionts: Bu lletin of Marine Science, v. 26, p. 530-537 Lee, J.J., Crocket, L.J. axonomic identity and physiological ecology of Chlamydomonas hedleyi sp. nov., algal flagellate symbiont fr v. 9, p. 407-422 Lee, J.J., Faber, W Endosymbiotic Symbiosis, v Lee, J.J anallock, P. (198gal symsi s as the driving force in the evolution of largoram J.J., Mer.ison, 198rimental studies of larger fora Foramnif Foramnif M J.J., Mer W.W.,Jr. and Lagziel, A. (1989) Identific ation and distribution of endosymbiotic diatoms i 35, p.353-366 Lee, J.J., Mles, J., Symon., Halloc k, P. (1995) Diatom symbionts in larger fora 105 Lees, R.P.,ns atis, The President, during growth-i : Journal of Experimenta Leutenegger, S. (1984) Symbiosis in be nthic foraminifera: Specificity and host Adations 4, p. 16-35 Leutenegge. and H19 radiot racer studies of pore function in foram Ha ge n, J. a nd St on e, R ( 19 74) Th e t o m the fo ram in ife r A rch ai as ang ul atu s: British Journal of Phycology, .W., Nathanson, B., Rottger R., Nishihira, M., and Kruger, R., (1993) ram di atom s fro m lar ge r fo i nifera collected in Pacific habitats. 1 4 (1 -3 ), p 7). 2 Al 65 -28 1 d H er f cEn minifera and their symbionts from the i i cEn ora minifera from Eva pta r, S bio inif era A nn N .Y A cda de my of Science, v. 503, p. 330-347 Lee Lee, J.J., McEnery, M.E., and Kahn, E.G., (1979) Lee y, M eral era: y, M n la E. a ese nd ar G ch, arr v. J.R ), p ( 3 0) Gulf of Elat on the Red Sea: Journal of 7. Ex pe R 10 (1 1-4 Sym p.1 biosis and the evolution of larger -14 ic rop ale on tol og y, v. 2 5, 18 0. E., ter K uile B ., Er ez, J., Rottger, R., Rockwell, R.F., Faber, rge r f ora mi nif era : M ic rop ale on tol og y, v s, A Caribbean hosts: Ma rine Micropalenotology, v. 26 (1-4), p. 99, E .H ., N nv ich itr ol o an s, J d .R sub (1 se 99 que 1) Photosynthesis in clem nt inv iv o a ccl imatization l B ota ny v 42 p p. 60 5-6 10 : Jo urn al of Fo ram in ife ral Re sea rc h, v 1 ans i en, nifera: Marine B H. J. ( 79 ) U ltr iology, v. 54, p. 11-16. astr uc ture a nd

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Lutze, G.F., and Wefer, G., (1980) Habitat and asexual reproduction of Cyclorbiculina compressa (Orbigmy), Soritidae: Journal of Foraminiferal Research, v. 10, p. 251260 Mallekh, R. and Lagerdere, J.P. (2002) Effect of temperature and dissolved oxygen concentration on the metabolic rate of the turbot and the relationship between metabolic scope Biology, v. 60 (5), p.11051115. McKee, E.D., Chr ntary belts in the lagoon of Kapingimara ciation of Petroleum Geologists, v. 43, p. 501-62 Muller, P. (1978) Carbo biont System: Journal of Foram Murry, J.W991) Ecology and distrtio inifera, in Biology of Foraifera demic Press, Inc., San Diep. scatine,19 logy of Coral Reefs. (eds A c., New York, pp.77-115 scatine,19 nd energy flux in reef corals: In Coral Reefs (ed. Dubinsky, Z.) Ecosyste orld, Elsevier, Amsterdam, p. 75-88. ly, M.B99atic,pirat ory and photosynthetic responses of Halodule wrightii to whole plant carbon budget. Thesis (M.S. Onuf, C.P. (1996) the Laguna Madre, Texas: Bul of M Richardson, B tion of unicellular algae to irradiance: An analysis of stra tegies: New Phytologist, v. 93, p.157-191 Rink S, Ku, Bij respion inversa: Marine Biology, v. 131 p. Rogers, C.S979) Th Puerto Rico: Limnology and Oceanograp 76 an d f eed in g d em an d: Jou rna l o f F ish on ic, ngi J., A an tol d L l: B eo ul pol leti d, n o E.B f t ( he 19 Am 59) er S ica ed n A ime sso n f ixa tio n a nd los s in a fo ra mineral-algal sym iniferal Research, v. 8 (1), p. 35-41. (1 min go, L. ( D. L. ( (1 letin K. hl M irat (4), ., (1 ibu n of benthic foram (e -25 ds. 4 Le e, J .J and A nd ers on O .R. ), A ca 221 73) Jo 90) 6) S lig Mu Mu Nee N ne utr s a itio nd n o R. f c En or de als an) in A B ca iolo dem gy ic an Pr d G ess eo In Th e rol e o f s ym bio tic al gae in m ca s of the W rbo n a om res ht )--U red ni uc ve tio rsit n i y o n T f S am ou pa th B Flo ay, rid in c a lu din g a B iom ari as ne s p Sc atte ien rn ce s in v. se 58 ag (2 ras ), p s m 4 ea 04 d o -42 ws 0 of ea rda ll, J ., and R ave n, J.A ., (19 83 ) A da pta ma J, (19 98 ) M ic r osensor studies of photosynthesis and th 3-5 e s 95 ymb iotic f o ram in ife r O rb uli na u ni 58 e p hy ro v duc 24 tiv (2 ity ), of p. 3 Sa 42 n C -3 ri 49. sto bel Re ef,

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77 Smith, D.F., (1977) Primary pro fera-zooxanthellae symbionts: Proceedings, Third International Coral Reef Symposium, Ro sentiel Scool of Marine and Atmospheric Scienc es, University of Miami. lanktonic foraminifer Orbulina universal contribution to ocean primary d Fo 15 s, J.F., Hershz, J.J. and Boutilier., (2000) Effects of ambient PO2 and temperature on oxygen uptake in Naus pompilius: Journal of Comparative Physiology B-Biochemical Systemic and Environmental Physiology, v. 170 (3), p. 231-236 H.K. (2002)logy of symbiont loss (bleaching) in Amphistegina gibbosa (class Foramra): Thesis (Ph.D.)versity of South F H.K. and H P., (2003) Ultrastr al responses in fieeached and experimentally stressed Amphistegina gibbosa (Class foramra): Journal of Eukaryotic Microbiology, v. 50 (5), p. 324-333 aylor, D., (1973) Symbiotic pathways of car bon in coral reef ecosy stems: Helgolander wiss. Meeresunters, v. 24, p. 276-283 ng in the metabolism of larger symbiont bearing foraminifera: Symbiosis, v. 4, p. 335-350 ter Kuile, B. (1991) Mechanismsalc ation and carbycli ng in algaring foraminifera, in Biology of Forara (eds J.J d O.R. And), Academic ew Yo 7 ofer, E.A. ais, J.L.6 vo microtuburing experimly induced conversions between tubulin assembly states in Allogromia lllCell Motili the Cyto34, p. 81-94 ms, D.E. and Hallock, P. (2004) Bleaching in Amphistegina gibbosa dg (Class Forara): obsom laboratory experiment s using visible and ultravioletOI: 10./-004-1351-5 r, L.W., (1e determioissolved oxyn water. Ber es, v. 21, 3-2846 Zar, J.H. (1984) Biostatistical An alysis, Prentice Hall, Inc. 718 p. ductivities of two foramini Spero H.J. and Parker, S.L., (1985) P hotosynthesis in the symbiotic p and its potentia pro uctivity: Journal of raminiferal Research, v. (4), p. 273-281 Staple kowit R.G tilu Talge, Cyto inife -U ni lorida 131 p. Talge, allock, uctur ld-bl inife T ter Kuile, B.,Erez, J., and Lee, J.J. (1987) Th e role of feedi for c i fic on c l-bea minife Lee an erson Press, N rk, p. 3-89 Welnh nd Trav (199 ) In vi le du ental atico aris: ty and skele ton, v. Willia Orbi ny minife ervati ns fro light D 1007 s00227 Winkle 888) Th inat n of d gen i Dtsch Chem G p. 284

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78 Zmiri, A., Kahan, D., Hochstein, S., and Reiss, and thermotaxis in some species of Ampistegina (Foraminifera): Journal of Protozoology, v.21, pp 133-138 Z., (1974) Phototaxis

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s 7 9 A ppe nd ice

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Appendix A Table A-1. Physical Characteristics of Archaias angulatus Specimen Mass Maj. Dia Inter. Diam Min. Diam. Surface Area (mg) (mm) (mm) (mm) (mm2) AA01 2.20 3.00 2.45 0.35 5.77 AA02 1.62 2.90 2.25 0.35 5.13 AA03 1.72 5.10 AA04 1.3.58 AA05 1.08 3.21 AA06 1.754.16 AA07 1.433.91 AA08 3.85 10.52 AA09 1.5.45 AA10 2.08 5.13 Mean 1.5.19 2.2.2. 95 40 15 2.1.1. 20 90 90 0.00. 35 .3555 06 2.2. 65 55 2.1. 00 95 00 .55.50 4.2. 25 95 3.2. 15 35 00 .55.55 79 2.2. 90 87 2.2. 25 24 0.0 55 .47 86 Median 1.74 2.90 2.23 0.53 5.11 Std. Dev02.06 .79 0. 56 0. 37 0.10 Table A-hlo SpecimChl a Mass g Chl a/mg foram Surface Area g Chl a/mm2 2. Cen rop hyll a E xtr act ion Arc hai as ang ula tus ( ug) (m g ) (m m2 ) AA01 0.0.09 50 2. 20 0. 23 5. 77 AA02 0.0.05 AA03 0.0.05 AA04 0.19 0.05 AA05 0.0.13 AA06 0.63 1.75 0.36 4.16 0.15 AA07 0.0.10 AA08 1.18 3.85 0.31 10.52 0.11 AA09 0.550.10 AA10 0.540.11 Mean 0.49 1.89 0.26 5.19 0.09 24 25 1.621.72 00 .15.14 55 .13.10 1.06 1.08 0.0 18 .39 3.3 58 .21 42 39 1.75 0 .22 3 .91 1.792.08 00 .31.26 55 .45.13 Median 0.0.10 Std. Dev0.29 0.04 46 1.750.78 00 .24.09 52 .11.06 80

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81dix AontinueTable A-3. Oxygen Production at experimental light intensities Archaiasulanmoles hr-1 ug Chl a-1 ) e (l photon Ap pen (C tus (mo d) angnsity Lig ht I nt m-2s1) Specimen 0 0.4.2 54 96 17 .4 5 36 .5 48 .4 92 .1 17 9 33 2 66 3 77 9 86 4 12 88 0 AA01 -21.15 -24.17 15.10 36.25 48.33 75.52 99.612 117.81 114.79 93.65 99.69 57.40 -84. AA02 -43.07 -55.38 6.15 55.38 67.68 86.14 110.11 129.22 116.91 92.30 116.91 92.30 -17AA03 -12.14 48.57 109.29 133.58 145.72 182.15 244 19 218.58 176.08 194.29 170.01 145.72 -109.29AA04 -23.71 23. 15 189.69 150.17 158.07 134.36 134.36 -11AA05 0.00 7.9 75 89.71 82.53 71.77 68.18 53.83 -50. AA06 -9.54 -14.7.23 69 69.15 83.46 90.61 81.07 76.30 -59.AA07 -3.87 -11..37 85 112.18 AA08 -8.89 -8.89 15.24 43.17 57.14 93.97 113.02 11 128.25 AA09 -5.42 -10.83 10.83 21.67 24.38 40.63 51.45 65.00 56.88 18.96 46.04 24.38 -70.AA10 -8.27 -2.76 13.79 22.06 38.60 49.63 82.79 113.05 110.29 110.29 93.75 93.75 -11Mean -13.61 -4.85 17.38 41.84 55.40 79.48 97.310 123.26 109.81 103.32 99.53 81.33 -95. 9 76 0.830.764.298.07.36.15.116.826.880.997.83 582.29 06.10.6.5 71 18 017 .00.9 234 35 .71.8 8 43 79 .04.0 16 53 02..8 75 13 64 5 8.5524612470425.8147 3161 4 -1 .779.3 4 1134 .92.82 1930 .08.95 40.69. 5463 5 7 7 1 85.121. 190 8 11 8.94.2 7 8 7710 .377.9 4 4292 .55.70 -81.-92. 6 2 9 Median -9.21 -9.86 12.31 35.35 45.70 72.58 91.210 115.43 112.54 92.97 96.72 84.30 -88. Std. Dev 12.67 27.642.43 48.61 35.11 47.30 35.37 38.68 35.48 0 1 0.87 64 75 3 4.0 7 34 .54 36 .87 42. 20 4 4.

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82 Ap pen di x A (C on tin ue d) Table A-4. O xyg en Productio n at exp eri men t al lig ht in ten s ities Arch ai as an gu lat us (n m oles hr -1 mm-2 ) L igh t In ten sity (u mol ph oton m -2s-1 ) Speci men 0 0. 96 1 7.4 5 3 6.5 48. 4 9 2.1 17 4.9 23 3 54 2 663 779 8 64 12 88 0 AA01 -1.82 -2.08 1.30 3.12 4.16 6.50 8.58 10.39 10.13 9.87 8.06 8.58 4.94 -7. AA02 -2.05 -2.63 0.29 2.63 3.22 4.10 5.27 5.27 6.15 5.56 4.39 5.56 4.39 -8.20 AA03 -0.59 2.35 5.30 6.47 7.06 8.83 10.01 9.42 10.59 8.53 9.42 8.24 7.06 -5. AA0 AA0 AA06 -1.44 -2.16 0.72 1.80 2.88 6.13 8.65 10.45 10.45 12.61 13.69 12.25 11.53 -9. AA07 -0.36 -1.08 -1.80 3.24 2.88 6.49 7.21 7.93 10.45 7.93 8.29 7.21 3.96 -7. AA08 AA0 AA10 -0.88 -0.29 1.46 2.34 4.10 5.27 8.78 9.66 12.00 11.71 11.71 9.95 9.95 -12. Mean -0.99 -0.58 1.24 3.26 4.30 6.45 8.07 9.02 10.25 9.44 8.81 8.46 6.89 -8.00 28 30285501 4 5 -10. .26 00 1.0. 2694 0.02. 034 1.4. 26 68 4.15. 961 5.7. 45 01 5.8. 86 42 9. 8.3 882 10.11.6 05 9 7.910.7 6 5 89. .38 35 8. 7 .12 88 7. 7. 12 01 -6.-6. 57411629 9 -1-0 .00.55 -1.-1. 0010 1.71.1 1 0 4. 2. 8520 6.42.4 2 8 10 4 .56.13 125. .7023 13.5.7 139 14.6.6 411 1 3.75.7 0 9 11 2.8.93 4 124 .13.68 102. .4148 -10.-7. Median -0.94 -1.04 1.20 2.88 4.13 6.31 8.50 9.54 10.45 9.20 8.86 8.41 7.04 -7. Std. 4208 Dev 0 .65 1. 63 1.8 2 1. 61 1.5 8 2 .01 2. 31 2.3 1 2.4 2 2.7 7 3 .61 2 .50 3. 01 2.

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83 d) ic scope and factorial metabolic scope for Archaias angulatus e h a ce orial Scope Appendix A (Continue Table A-5. Metabol Specim n N ormalized to C l a Norm lized to surfa area Fact AA01 4.0 63.4 5.5 AA0 2 4.0 3 9.0 4 5.0 5 1.0 6 6.3 7 21.0 8 10.4 9 13.0 0 14.0 9.6 129.2 6.2 AA0 97.2 4.7 AA0 94.8 5.0 AA0 50.2 6.6 AA0 50.1 7.6 AA0 77.4 7.2 AA0 83.8 9.4 AA0 65.0 6.6 AA1 107.5 11.4 Mean 81.9 7.0 Medi an 9.0 ev 5.7 80.6 6.6 Std. D 25.7 2.1 Table A-6ved pters fromnthesisance curv Normalized to Chl a Normalized to surface area Deri arame Photosy /Irradi es Specimen P AlphaI PmaxAlphaIk max k AA01 129.40 1.52 84.91 11.13 0.13 84.90 AA02 155.20 2.60 59.72 7.38 0.12 59.73 2 3 0.28 35.21 AA04 177.70 1.75 101.5 9.42 0.10 96.80 5 .77 0.89 84.79 9 0.12 84.90 6 .15 0.55 163.05 1 0.08 162.96 7 .33 0.81 116.67 8 0.08 116.66 8 7.10 1.47 86.58 1 0.17 86.60 9 .76 0.72 75.78 5 0.07 75.69 0 3.80 0.77 147.72 1 0.08 147.71 22.05 1.68 95.60 1 0.12 95.12 AA03 20 .30 5.74 5.26 9.80 4 AA0 75 .87 AA0 90 3.62 AA0 94 .79 AA0 12 4.28 AA0 54 .57 AA1 11 2.08 Mean 1 0.19 Medi an 0.45 1.18 85.74 9 0.11 85.75 v. .24 1.56 38.55 2 0.06 38.49 12 .84 Std. De 46 .69

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84 Table B-1. Physical Characteristics of Cyclorbiculina compressa ecimeaDia er. Diam MinSurface Aa Appendix B Sp n M ss Maj. Int Diam. re (mm) m) (m (mm (m g) (m m) 2) CC01 3.853.6 9 9 2.9 10.8 2. CC02 3.953.65 2 .1 5.35.1 3 3 3.53.45 1 3.63.35 3 5.65.5 9 7 54.75 5 .2 3.83.5 5 7 4.154.15 3 .3 3.63.25 1 34.244.03 4 7 3.1 11.3 3 CC03 6.3 21.2 6. CC04 2.1 9.48 2. CC05 2.3 9.47 2. CC06 7.7 24.1 7. CC07 5.2 18.6 5 CC08 2.7 10.4 2. CC09 3.3 13.5 3 CC10 2.1 9.19 2. Mean .77 13.8 3.7 Medi an 3.93.63.1 3 1.0.77 0.81 3 11 Std Dev 95 5.49 1.95 Table B -2.r a Extract ssa eh g/mg foram ce Area 2 Chlo ophyll ion Cyclorbiculina c ompre Specim n C l a Mass Chl a Surfa g Chl a/mm g) m2) (u g) (m (m CC01 0. 290 08 .89 02 4 2. 0. 10 0. CC02 0. 210 08 .32 02 630 11 .23 03 110 09 48 02 130 08 47 02 870 10 .19 03 620 13 .65 04 270 08 .45 02 430 14 .53 03 210 10 19 02 377 10 .84 03 5 3. 0. 11 0. CC03 0. 9 6. 0. 21 0. CC04 0. 9 2. 0. 9. 0. CC05 0. 8 2. 0. 9. 0. CC06 0. 0 7. 0. 24 0. CC07 0. 5 5. 0. 18 0. CC08 0. 3 2. 0. 10 0. CC09 0. 5 3. 0. 13 0. CC10 0. 0 2. 0. 9. 0. Mean 0. 9 3. 0. 13 0. Medi an .200 09 .11 02 v. 295 02 49 01 0 4 3. 0. 11 0. Std. De 0. 4 1. 0. 5. 0.

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Appendix B (Continued) Table B-3. Oxygen Production at experimental light intensities Cyclorbiculina compressa (nmoles hr-1 ug Chl a -1 ) -1) Light Intensity (umol photon m-2s Specimen 06.48.4880 0.9 17 5 36 .54 9217 .1 4.9 233 542 663 779 864 12 CC01 -17.22-1 -28.48 5.65 28.6 945.9 1.6.311 -124.36 6.20.21551-177.29 60.2911101-137.32111.8.4891N1 -229.43 90.88N31-316.45 837.3N11-106.91 30.26.1.14131-126.48 37.585.8171.19N.191-225.05 8.8.214N1-124.470.9.21099 -246.52 6.2.112121-181.43 62 5 86 07 9 .22 3.90 N.A. 9 1.81 5. 80.34 74.60 CC02 16.1 -16.11 15.7 21.0 131.2 9781. 1 115 301 5.14.5 0.13 N.A. 1N.A. 30 36.56 109.50 CC03 -32.86 -36.45 -1.90 7.67 26.68 5 74 7 8 1 1.27 8.231 01.68 90.84 CC04 -54.82 -63.03 -13.8 20.4 27. 54 2 76 8 .09 1 0.44 .A. 1.A. .24 75.38 48.32 CC05 -14.28 -22.06 -7.25 7.19 14.4 79.1 1 10 698 .88 6.94 1 7.65 09.47 14.49 CC06 -6.58 -10.02 29.1 52.6 66.2 94.7 0 96 4 124813 0.1816 25.73 .A. 1N.A. 5.98 06.13 87.81 CC07 -2.03 2.01 14.1 8 31.9.2 61.24 97 0 121 3 7.83 8.72 46.75 98.50 CC08 0.00 0.00 29.1 8 64 076. 11 1 7 75 9.88 A. A. 5.44 88.12 135.21 CC09 -14.37 -8.75 11.5 23.5 032.3 186 4 94 00 1 6.32 5.80 .1N.A. 43.94 17.51596 78.73 CC10 -40.27 -46.99 -19.8 4 13.3 319.7 59 6 79 36 9 .59 0.69 .21 .9 59.09 Mean -19.85 -22.99 6.27 27.0 40.2 178 1 95 99 1 2.83 9.26 N.A. 9.75 12.19 79.71 Median 80.6.11011N.121-157.30 .3.73N 69.69 -15.24 -19. 0 8.62 22.2 631. 8 7 8 6 95 7 6.7 0 9.81 A A. 6.81 07.80 83.27 Std. Dev 17.57 20.98 16.88 18.6 021.1 521 9 17 59 2 .26 2.62 .3 1.06 37.76 33.67 85

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86 les h-2) -1 Appendix B (Continued) able B-4. Oxygen Production at experimental light intensities Cyclorbiculina compressa (nmor-1 mm T Light Inty (upho2s ensit mol ton m) Specim e n 5 1 54667948 0 0.96 17.4 36. 5 48.4 92.1 74.9 233 2 3 7 86 128 0 CC01 8 2.N.A.053 6 3.N.42 76 3..50 87 2..29 74 1..60 2 4.N.80 7 5.N.84 4.N.A.251 8 4.N.A.844 83 2.N.A118 3.N.36 -0.3 -0.62 0.12 0.63 1.00 1.37 1.88 2.04 49 2 0 1.7 1.6 -2.71 CC02 -0.3 -0.36 0.35 0.46 0.69 1.73 2.44 2.76 32 A. 3. 3 3.0 2.42 -3.92 CC03 -1.0 -1.18 -0.0 0.25 0.87 1.66 2.41 2.91 62 N A. 3. 2 3.3 2.95 -4.46 CC04 -1.0 -1.25 -0.2 0.40 0.55 1.08 1.51 1.76 18 N A. 2. 0 1.4 0.96 -4.54 CC05 -0.2 -0.42 -0.1 0.14 0.28 1.51 1.93 1.70 3 66 N A. 2. 4 2.1 0.28 -6.06 CC06 -0.2 -0.33 0.96 1.74 2.19 3.13 3.18 4 .97 15 A. 3. 3 3.5 2.90 -3.53 CC07 -0.0 0.07 0.50 1.12 2.14 3.41 .43 4.61 18 A. 4. 6 5.1 3.45 -4.43 CC08 0.00 0.00 0.63 1.38 1.65 2.53 2.50 3.70 31 4. 1 4.0 2.9 -4.85 CC09 -0.4 -0.29 0.39 0.79 1.08 2.91 3.15 4.24 89 4. 3 3.9 2.6 -4.18 CC10 -0.8 -1.02 -0.4 0.29 0.43 1.31 1.73 2.02 19 2. 6 1.3 1.29 -5.37 Mean -0.4 -0.54 0.20 0.72 1.09 2.06 2.52 2.97 40 A. 3. 7 2.9 2.14 -4.40 Median Std. De -0.30. 794 3.N.A.463 v 40 47 44 53 69 85 0.88 1.1.N.04 93 -0.30. 0.20. 0.550. 0.930. 1.690. 2.43 2.84 09 47 23 3A. 1. 7 3.108 1. 2.527 1. -4.450.

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87 a Appendix B (Continued) Table B-5. Metabolic scope and factorial scope for Cyclorbiculina compress Specimen Normalized to Chl a Norma lized to surface area Factorial scope CC01 1 07.14 2.34 7.22 CC02 1 1 1 4 1 4 3 2 1 1 1 6 2 1 1 8 2 6 1 1 61.18 3.56 1.00 CC03 04.47 3.39 .18 CC04 74.60 3.45 .18 CC05 02.17 5.79 2.16 CC06 00.33 3.31 6.25 CC07 24.46 4.36 2.46 CC08 25.05 4.85 .00 CC09 10.11 3.69 .66 CC10 06.24 4.49 .12 Mean 61.57 3.92 5.81 Median 1 8 1 42.82 3.63 .66 Std Dev. 66.64 0.97 8.47 Table B-6. Derived parameters from Photosynthesis/Irradiance curves Normalized to Chl a Normalized to surface area Specimen P max Alpha I k P max Alpha I k CC01 115.20 1.24 93.28 2 2.51 0.03 93.2 CC02 165.90 1.04 160.14 3.67 4 5 0 6 4 8 3 5 7 0 0.02 160.0 CC03 136.30 1.07 127.03 4.43 0.04 127.1 CC04 152.70 1.83 83.40 3.02 0.04 83.5 CC05 124.90 0.92 136.12 2.39 0.02 136.1 CC06 121.60 1.72 70.82 4.02 0.06 70.8 CC07 143.50 1.23 116.57 5.03 0.04 116.5 CC08 188.60 1.29 146.54 4.07 0.03 146.7 CC09 155.50 1.03 150.39 5.22 0.04 150.3 CC10 136.80 1.35 101.48 2.98 0.03 101.4 Mean 144.10 1.27 118.58 3.73 0.03 118.6 Median 140.15 1.23 121.80 3.84 6 7 0.03 121.8 Std. Dev. 22.32 0.30 30.47 1.00 0.01 30.4

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88 m ea Appendix C Table C-1. Physical Characteristics of Amphistegina gibbosa Specimen Mass Maj. Diam. Inter. Diam Min. Dia Surface Ar (mg) (mm) ) 2) (mm) (mm (mm AG01-1 0.27 0.90 0 64 0.90 0.4 0. AG01-2 0.23 0.90 5 60 5 57 5 71 0 90 5 67 0 57 0 12 0 67 0 99 0 64 5 78 0 99 0 86 5 37 1 0 75 0 90 0 82 G04-4 0.60 1.30 1.15 0.50 1.17 G04-5 0.65 1.35 1.10 0.55 1.17 G05-1 0.27 1.00 0.85 0.35 0.67 0 0.82 G05-3 0.42 1.10 1.00 0.45 0.86 A 0 AG05 00 0 0.86 5 .750 0.50 0 0.75 0.30 0.47 0 1.00 0.45 0.79 5 0.95 0.40 0.78 5 1.00 0.50 0.90 5 0.65 0.25 0.38 5 0.75 0.35 0.50 5 0.85 0.40 0.63 0 0.90 0.40 0.78 5 1.10 0.50 1.08 5 0.65 0.30 0.38 0. 75 0.35 0.47 AG08-3 0.24 1.00 0.85 0.40 0.67 0.85 0.3 0. AG01-3 0.29 0.90 0.80 0.4 0. AG01-4 0.28 1.00 0.90 0.3 0. AG01-5 0.44 1.15 1.00 0.5 0. AG02-1 0.36 0.95 0.90 0.4 0. AG02-2 0.27 0.90 0.80 0.4 0. AG02-3 0.47 1.30 1.10 0.5 1. AG02-4 0.33 1.00 0.85 0.4 0. AG02-5 0.52 1.20 1.05 0.5 0. AG03-1 0.23 0.90 0.90 0.4 0. AG03-2 0.36 1.05 0.95 0.3 0. AG03-3 0.60 1.20 1.05 0.5 0. AG03-4 0.49 1.10 1.00 0.4 0. AG03-5 0.83 1.40 1.25 0.6 1. AG040.20 1.00 0.95 0.4 0. AG04-2 0.38 1.15 1.00 0.5 0. AG04-3 0.34 1.10 0.95 0.4 0. A A A AG05-2 0.42 1.10 0.95 0.4 A G05-4 0.51 1.1 1.05 0.50 0.91 -5 0.40 0. 1.10 1. .45 .35 AG06-1 24 0.8 0 AG06-2 0.24 0.8 AG06-3 0.34 1.0 AG06-4 0.29 1.0 AG06-5 0.38 1.1 AG07-1 0.10 0. 0.7 AG07-2 12 0.8 AG07-3 0.16 0.9 AG07-4 0.30 1.1 AG07-5 0.53 1.2 AG08-1 AG08-2 0.10 10 0.7 0.80 0.

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89 ) Speen iam. In. Diam. MinSurface Aa Appendix C (Continued) Table C-1 (Continued cim Mass Maj. D ter Diam. re (mg) (mm ) (mm m) (mm2) ) (m AG08-3 0.24 1.00 0. 850.40 0.67 AG08-4 0 850.40 0.73 950.45 0.82 1 750.30 0.50 0 800.40 0.57 050.50 0.99 0 000.55 0.94 AG09-5 0.58 1.35 1.15 0.55 1.22 AG10-1 0.15 0.80 0.75 0.30 0.47 AG10-2 0.16 0.80 0.75 0.35 0.47 AG10-3 0.21 0.85 0.75 0.35 0.50 AG10-4 0.22 0.95 0.85 0.40 0.63 AG10-5 0.40 1.25 1.05 0.45 1.03 Mean 0.34 1.04 0.92 0.42 0.76 .37 1.10 0. AG08-5 0.40 1.10 0. AG090.14 0.85 0. AG09-2 .20 0.90 0. AG09-3 AG09-4 0.54 .45 1.20 1. 1.20 1. Median 0.34 1.03 0.93 0.40 0.76 Std. Dev. 0.16 0.17 0.14 0.08 0.23 Table C-2. Chlorophyll a Extraction Amphistegina gibbosa Specimen Chl a Mass ug Chl a/mg foram Surface Area g Chl a/mm2 (ug) (mg) (mm2) AG01-1 0.07 0.27 0.26 0.64 0.11 AG01-2 0.04 0.23 0.19 0.60 0.07 AG01-3 0.04 0.29 0.13 0.57 0.07 AG01-4 0.05 0.28 0.19 0.71 0.08 AG01-5 0.07 0.44 0.16 0.90 0.08 AG02-1 0.06 0.36 0.18 0.67 0.09 AG02-2 0.04 0.27 0.14 0.57 0.06 AG02-3 0.12 0.47 0.25 1.12 0.10 AG02-4 0.06 0.33 0.17 0.67 0.09 AG02-5 0.08 0.52 0.15 0.99 0.08 AG03-1 0.03 0.23 0.13 0.64 0.05 AG03-2 0.09 0.36 0.24 0.78 0.11 AG03-3 0.08 0.60 0.13 0.99 0.08 AG03-4 0.10 0.49 0.20 0.86 0.11

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90 Appendix C (Continued) Table C-2 (Continued) Specimen Chl a Mass ug Chl a/mg foram Surface Area g Chl a/mm2 (ug) (mg) (mm2) AG03-5 0.14 0.83 0.17 1.37 0.10 AG04-1 0.05 0.20 0.27 0.75 0.07 AG04-2 0.07 0.38 0.18 0.90 0.07 AG04-3 0.10 0.34 0.29 0.82 0.12 AG04-4 0.11 0.60 0.18 1.17 0.09 AG04-5 0.12 0.65 0.19 1.17 0.10 AG05-1 0.05 0.27 0.19 0.67 0.08 AG05-2 0.06 42 0.14 0.82 0.07 AG05-3 0.23 0.86 0.11 AG05-4 0.51 0.19 0.91 0.11 AG05-5 0.23 0.86 0.11 AG06-1 0.11 0.50 0.05 6-2 24 0.12 0.47 0.06 6-3 0.22 0.79 0.09 6-4 29 0.17 0.78 0.06 6-5 0.28 0.90 0.12 0.26 0.38 0.07 7-2 0.12 0.33 0.50 0.08 7-3 0.25 0.63 0.06 7-4 30 0.32 0.78 0.13 7-5 0.17 1.08 0.08 AG08-1 0.27 0.38 0.07 AG08-2 0.10 0.34 0.47 0.07 AG08-3 0.19 0.67 0.07 AG08-4 0.12 0.73 0.06 AG08-5 0.40 0.13 0.82 0.06 AG09-1 0.14 0.23 0.50 0.06 AG09-2 0.20 0.25 0.57 0.09 AG09-3 0.54 0.20 0.99 0.11 AG09-4 0.45 0.30 0.94 0.14 AG09-5 0.58 0.20 1.22 0.10 AG10-1 0.15 0.24 0.47 0.08 AG10-2 0.16 0.25 0.47 0.09 AG10-3 0.21 0.22 0.50 0.09 AG10-4 0.22 0.35 0.63 0.12 AG10-5 0.40 0.19 1.03 0.07 Mean 0.34 0.21 0.77 0.09 0. 0.42 0.10 0.10 0.09 0.03 0.03 0.07 0.05 0.11 0.03 0.04 0.04 0.10 0.09 0.03 0.03 0.05 0.04 0.05 0.03 0.05 0.11 0.14 0.12 0.04 0.04 0.05 0.08 0.07 0.07 0.40 0.24 AG0 AG0 AG0 AG0 AG0 AG0 AG0 AG0 AG0 0. 0.34 0. 0.38 0.10 71 0.16 0. 0.53 0.10 0.24 0.37 Median 0.34 0.20 0.78 0.08 Std. Dev. 0.16 0.06 0.24 0.02 0.06 0.0 3

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91 Appendix C (Continued) Table C-3. Oxygen Production at experimental light intensities Amphistegina gibbosa (nmoles hr-1 ug Chl a-1 ) Light Intensity (umol photon m-2s-1) Specimen 0 0.96 3.13 10.91 17.4536.5 48.4 92.1 174.9233 542 779 1288 0 AG01 -5.48 -5.48 5.478 5.478 16.4416.440 27.3910.9610.9621.91 21.915.478-76.7 AG02 -17 -12.8 -4.26 12.77 17.0221.2817.0221.2821.2812.7721.28 0 -12.8 -55.32 AG03 -27.6 -27.6 -24.1 -6.89 -6.89 0 0 -3.45 0 -6.89 0 -3.45 -34.5 -55.14 AG04 -36.8 -40.2 -26.8 -16.7 -6.7 0 -3.35 -6.7 -3.35 -6.7 -3.35 -10 -36.8 -63.6 AG05 -68.6 -64.8 -57.1 -45.7 -38.1 -26.7 -26.7 -26.7 -26.7 -26.7 -26.7 -26.7 -49.5 -80.01 AG06 -69.6 -64.3 -64.3 -53.6 -42.9 -26.8 -26.8 -21.4 -21.4 -26.8 -21.4 -26.8 -53.6 -91.07 AG07 -61.2 -71.4 -40.8 -20.4 -15.3 -5.1 -5.1 0 0 -5.1 0 -10.2 -40.8 -66.33 AG08 -14.9 -29.8 -14.9 0 7.4390 0 0 0 -7.44 0 -7.44 -52.1 -104.1 AG09 -10.1 -10.1 -6.75 10.12 16.8616.8616.8616.8616.8610.1213.49 6.745-13.5 -43.84 AG10 -27.3 -38.3 -32.8 5.465 21.8621.8627.3327.3327.3327.3338.26 16.4 0 -49.19 Mean -33.9 -36.5 -26.6 -10.9 -3.02 1.788-0.07 3.4622.498-1.84 4.35 -3.95 -28.8 -68.53 Median -27.4 -34 -25.5 -3.45 0.3720 0 0 0 -5.9 0 -5.44 -35.6 -64.96 Std. Dev 24.35 23.93 22.75 23.17 23.3417.9717.6 19.2717.3917.3219.92 16.1221.9119.267

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Appendix C (Continued) Table C-4. Oxygen Production at experimental light intensities Amphistegina gibbosa (nmoles hr-1 mm-2) Light Intensity (umol photon m-2s-1) Specimen 0 0.96 3.13 10.91 17.45 36.5 48.4 92.1 174.9 233 542 779 1288 0 AG01 -0.44 -0.44 0.44 0.44 1.32 1.32 0.00 2.20 0.88 0.88 1.76 1.76 0.44 -6.15 AG02 -1.49 -1.12 -0.37 1.12 1.49 1.87 1.49 1.87 1.87 1.12 1.87 0.00 -1.12 -4.85 AG03 -2.58 -2.58 -2.26 -0.65 -0.65 0.00 0.00 -0.32 0.00 -0.65 0.00 -0.32 -3.23 -5.16 AG04 -3.43 -3.74 -2.50 -1.56 -0.62 0.00 -0.31 -0.62 -0.31 -0.62 -0.31 -0.94 -3.43 -5.92 AG05 -6.55 -6.18 -5.46 -4.37 -3.64 -2.55 -2.55 -2.55 -2.55 -2.55 -2.55 -2.55 -4.73 -7.64 AG06 -5.66 -5.23 -5.23 -4.36 -3.48 -2.18 -2.18 -1.74 -1.74 -2.18 -1.74 -2.18 -4.36 -7.40 AG07 -5.33 -6.22 -3.56 -1.78 -1.33 -0.44 -0.44 0.00 0.00 -0.44 0.00 -0.89 -3.56 -5.78 AG08 -0.98 -1.95 -0.98 0.00 0.49 0.00 0.00 0.00 0.00 -0.49 0.00 -0.49 -3.41 -6.83 AG09 -1.07 -1.07 -0.71 1.07 1.78 1.78 1.78 1.78 1.78 1.07 1.42 0.71 -1.42 -4.62 AG10 -2.41 -3.38 -2.90 0.48 1.93 1.93 2.41 2.41 2.41 2.41 3.38 1.45 0.00 -4.34 Mean -2.99 -3.19 -2.35 -0.96 -0.27 0.17 0.02 0.30 0.23 -0.15 0.38 -0.34 -2.48 -5.87 Median -2.50 -2.98 -2.38 -0.32 -0.07 0.00 0.00 0.00 0.00 -0.47 0.00 -0.41 -3.32 -5.85 Std. Dev 2.17 2.13 2.00 2.05 2.06 1.61 1.59 1.71 1.58 1.54 1.77 1.40 1.82 1.15 92

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93 Appendix C (Continued) Table C-5. Metabolic Scope and factorial scope for Amphistegina gibbossa Specimen Normalized to Chl a Norma lized to surface area Factorial Scope AG AG AG AG AG AG AG AG AG AG M M St Tabl 01 71.2 5.7 14.0 02 38.3 3.4 3.3 03 27.6 2.6 2.0 04 26.8 2.5 1.7 05 11.4 1.1 1.2 06 21.4 1.7 1.3 07 5.1 0.4 1.1 08 89.3 5.9 7.0 09 33.7 3.6 4.3 10 21.9 1.9 1.8 ean 34.7 2.9 3.8 edian 27.2 2.5 1.9 d. Dev. 26.2 1.8 4.0 e C-6. Derived parameters from Photosynthesis/Irradiance curves Normalized to Chl a Normalized to surface area Specimen PmaxAlphaIkPmaxAlpha Ik AG01 20.20 1.91 10.58 1.62 0.15 10.57 AG02 36.00 4.02 8.95 3.17 0.35 8.96 AG03 25.80 2.01 12.87 2.42 0.19 12.86 AG04 33.01 2.46 13.44 3.08 0.23 13.45 AG05 42.04 2.42 17.37 4.01 0.23 17.38 AG06 46.88 1.72 27.19 3.81 0.14 27.19 AG07 58.55 4.26 13.73 5.10 0.37 13.74 AG08 14.83 1.50 9.92 0.97 0.10 9.76 AG09 25.72 2.33 11.05 2.71 0.25 11.05 AG10 56.57 3.11 18.20 5.00 0.28 18.20 Mean 35.96 2.57 14.33 3.19 0.23 14.32 Median 34.51 2.37 13.15 3.12 0.23 13.15 Std. Dev. 14.92 0.94 5.44 1.34 0.09 5.45

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94 Appendix D Table D-1. Physical Characteristics of Amphistegina lessonii Specimen Mass Maj. Diam Inter. Diam Min Diam Surface Area (mg) (mm) (mm) (mm) (mm2) AL01-1 0.57 1.20 1.10 0.55 1.04 AL01-2 1.03 1.50 1.30 0.65 1.53 AL01-3 0.97 1.40 1.35 0.65 1.48 AL02-1 0.31 1.00 0.90 0.50 0.71 AL02-2 0.65 1.25 1.00 0.65 0.98 AL02-3 0.56 1.15 1.05 0.65 0.95 AL03-1 0.26 1.00 0.95 0.45 0.75 AL03-2 0.71 1.40 1.15 0.65 1.26 AL03-3 0.83 1.45 1.25 0.70 1.42 AL04-1 0.22 1.00 0.90 0.45 0.71 AL04-2 0.30 1.15 0.95 0.50 0.86 AL04-3 0.51 1.25 1.10 0.55 1.08 AL05-1 0.29 1.15 1.05 0.55 0.95 AL05-2 0.38 1.20 1.00 0.55 0.94 AL05-3 0.39 1.25 1.05 0.60 1.03 Mean 0.53 1.22 1.07 0.58 1.05 Median 0.51 1.20 1.05 0.55 0.98 Std. Dev. 0.26 0.16 0.14 0.08 0.27 Table D-2. Chlorophyll a Extraction Amphistegina lessonii Specimen Chl a Mass ug Chl a/mg foram Surface Area g Chl a/mm2 (ug) (mg) (mm2) AL01-1 0.136 0.57 0.2381.0370.131 AL01-2 0.228 1.03 0.2221.5320.149 AL01-3 0.205 0.97 0.2121.4840.138 AL02-1 0.103 0.31 0.3330.7070.146 AL02-2 0.144 0.65 0.2220.9820.147 AL02-3 0.118 0.56 0.2110.9480.124 AL03-1 0.08 0.26 0.3090.7460.107 AL03-2 0.137 0.71 0.1921.2640.108 AL03-3 0.194 0.83 0.2341.4240.136 AL04-1 0.072 0.22 0.3260.7070.102 AL04-2 0.076 0.3 0.2540.8580.089

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95 Appendix D (Continued) Table 2 (Continued) Specimen Chl a Mass ug Chl a/mg foram Surface Area g Chl a/mm2 (ug) (mg) (mm2) AL04-3 0.111 0.51 0.218 1.080.103 AL05-1 0.116 0.29 0.3990.9480.122 AL05-2 0.12 0.38 0.3160.9420.127 AL05-3 0.135 0.39 0.3461.0310.131 Mean 0.132 0.53 0.2691.0460.124 Median 0.12 0.51 0.2380.9820.127 Std. Dev. 0.046 0.26 0.0630.2680.018

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96 Appendix D (Continued) Table D-3. Oxygen Production at experimental light intensities Amphistegina lessonii (nmoles hrP P-1 P P P Pug Chl aP P-1 P P P P) Light Intensity (umol photon mP P-2P PsP P-1P P) Specimen 0 0.96 3.13 10.91 17.45 36.5 48.4 92.1 174.9 233 542 779 1288 0 AL01 -13.19 -13.19 -7.92 5.26 10.53 28.97 28.97 34.24 28.97 31.60 36.87 31.60 10.53 -44.80 AL02 -4.12 -4.12 -0.01 16.40 20.50 36.91 36.91 32.81 36.91 32.81 45.12 36.91 20.50 -73.87 AL03 -18.27 -21.92 -10.97 -0.01 10.94 21.89 18.24 25.54 25.54 18.24 29.19 21.89 -3.66 -62.07 AL04 -28.94 -40.50 -17.37 -5.80 5.77 23.12 23.12 28.91 40.48 28.91 46.26 23.12 -5.80 -104.13 AL05 -32.35 -32.35 -28.31 -20.23 -8.10 4.03 4.03 4.03 12.11 4.03 8.07 4.03 -24.27 -88.95 Mean -19.37 -22.42 -12.91 -0.88 7.93 22.99 22.26 25.11 28.80 23.12 33.10 23.51 -0.54 -74.76 Median -18.27 -21.92 -10.97 -0.01 10.53 23.12 23.12 28.91 28.97 28.91 36.87 23.12 -3.66 -73.87 Std. Dev. 11.53 14.55 10.63 13.56 10.43 12.15 12.34 12.26 11.08 12.12 15.60 12.52 17.07 23.04 Table D-4. Oxygen Production at experimental light intensities Amphistegina lessonii (nmoles hrP P-1 P PmmP P-2P P) Light Intensity (umol photon mP P-2P PsP P-1P P) Specimen 0 0.96 3.13 10.91 17.45 36.5 48.4 92.1 174.9 233 542 779 1288 0 AL01 -0.57 -0.57 0.00 2.28 2.84 5.12 5.12 4.55 5.12 4.55 6.26 5.12 2.84 -10.24 AL02 -2.18 -2.62 -1.31 0.00 1.31 2.62 2.18 3.06 3.06 2.18 3.49 2.62 -0.44 -7.43 AL03 -2.84 -3.97 -1.70 -0.57 0.57 2.27 2.27 2.84 3.97 2.84 4.54 2.27 -0.57 -10.21 AL04 -4.11 -4.11 -3.59 -2.57 -1.03 0.51 0.51 0.51 1.54 0.51 1.03 0.51 -3.08 -11.30 AL05 -2.31 -2.62 -1.54 -0.02 1.04 2.92 2.83 3.15 3.55 2.91 4.10 2.99 0.05 -9.09 Mean -2.18 -2.62 -1.31 0.00 1.31 2.62 2.27 3.06 3.97 2.84 4.54 2.62 -0.44 -10.21 Median 1.30 1.49 1.31 1.78 1.42 1.77 1.80 1.72 1.34 1.68 1.99 1.83 2.25 2.12 Std. Dev. -1.85 -1.85 -1.11 0.74 1.48 4.07 4.07 4.81 4.07 4.44 5.18 4.44 1.48 -6.29

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97 Appendix D (Continued) Table D-5. Metabolic Scope and factorial scope for Amphistegina lessonii Specimen Normalized to Chl a Norma lized to surface area Factorial Scope AL01 31.6 4.4 3.4 AL02 69.8 9.7 18.0 AL03 43.8 5.2 3.4 AL04 75.2 7.4 3.6 AL05 56.6 7.2 2.8 Mean 55.4 6.8 6.2 Median 56.6 7.2 3.4 Std. Dev. 18.0 2.0 6.6 Table D-6. Derived parameters from Photosynthesis/Irradiance curves Normalized to Chl a Normalized to surface area Specimen PBmaxB AlphaIk PBmaxB Alpha Ik AL01 45.90 1.67 27.50 6.48 0.23 27.76 AL02 41.45 1.85 22.43 5.79 0.26 22.61 AL03 41.81 1.92 21.82 5.04 0.23 22.16 AL04 61.84 2.09 29.59 6.31 0.20 32.17 AL05 38.98 1.48 26.39 5.02 0.19 27.09 Mean 46.00 1.80 25.55 5.73 0.22 26.36 Median 41.81 1.85 26.39 5.79 0.23 27.09 Std. Dev. 9.20 0.24 3.33 0.68 0.03 4.12

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98 Appendix E Table E-1. Physical Characteristics of Amphistegina radiata Specimen Mass Maj. Dia Inter. Diam Min. Diam.Surface Area (mg) (mm) (mm) (mm) (mmP P2P P) AR01 1.73 1.90 1.80 0.70 2.69 AR02 1.53 1.75 1.65 0.65 2.27 AR03 1.51 1.80 1.70 0.70 2.40 AR04 1.41 1.80 1.60 0.70 2.26 AR05 0.98 1.70 1.60 0.60 2.14 AR06 1.54 1.75 1.65 0.65 2.27 AR07 1.56 1.90 1.75 0.65 2.61 AR08 1.53 1.85 1.70 0.65 2.47 AR09 2.12 2.15 2.00 0.75 3.38 AR10 1.75 1.80 1.75 0.80 2.47 Mean 1.57 1.84 1.72 0.69 2.50 Median 1.54 1.80 1.70 0.68 2.44 Std. Dev. 0.29 0.13 0.12 0.06 0.35 Table E-2. Chlorophyll a Extraction Amphistegina radiata Specimen Chl a Mass g Chl a/mg foram Surface Area g Chl a/mmP P2P P (ug) (mg) (mmP P2P P) AR01 0.29 1.73 0.17 2.69 0.11 AR02 0.26 1.53 0.17 2.27 0.11 AR03 0.24 1.51 0.16 2.40 0.10 AR04 0.32 1.41 0.23 2.26 0.14 AR05 0.20 0.98 0.20 2.14 0.09 AR06 0.32 1.54 0.21 2.27 0.14 AR07 0.35 1.56 0.22 2.61 0.13 AR08 0.28 1.53 0.18 2.47 0.11 AR09 0.59 2.12 0.28 3.38 0.18 AR10 0.40 1.75 0.23 2.47 0.16 Mean 0.32 1.57 0.20 2.50 0.13 Median 0.31 1.54 0.20 2.44 0.12 Std. Dev. 0.11 0.29 0.04 0.35 0.03

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99 Appendix E (Continued) Table E-3. Oxygen Production at experimental light intensities Amphistegina radiata (nmoles hrP P-1 P Pug Chl aP P-1 P P) Light Intensity (umol photon mP P-2P PsP P-1P P) Specimen 0 0.96 3.13 10.91 17.45 36.5 48.4 92.1 174.9 233 542 779 1288 0 AR01 -25.68 20.55 20.55 -5.14 20.55 5.14 10.27 10.27 10.27 35.95 10.27 15.41 -30.82 -51.36 AR02 -34.82 -40.62 -17.41 -5.80 -11.61 0.00 -11.61 0.00 5.80 0.00 23.21 17.41 -29.01 -92.84 AR03 -18.46 -36.92 -24.61 -18.46 -18.46 -18.46 -18.46 -6.15 -18.46 -12.31 -6.15 -12.31 -24.61 -86.14 AR04 -9.31 -9.31 -4.65 9.31 0.00 13.96 4.65 23.27 23.27 13.96 41.89 41.89 46.54 0.00 AR05 -38.36 -92.07 -69.05 -61.38 -53.71 -53.71 -53.71 -53.71 -46.03 -61.38 -38.36 -53.71 -38.36 -130.43 AR06 -70.89 4.73 -80.34 -33.08 -51.98 -33.08 -23.63 -42.53 -42.53 -51.98 -66.16 -70.89 -51.98 -132.32 AR07 -42.84 -55.69 -42.84 -47.13 -34.27 -29.99 -38.56 -34.27 -29.99 -34.27 -8.57 -38.56 -38.56 -14.42 AR08 -37.86 -43.27 -32.45 -10.82 -21.64 -16.23 -16.23 -16.23 -16.23 -21.64 -5.41 -10.82 -37.86 -97.36 AR09 -20.35 -17.80 -15.26 -7.63 -7.63 -7.63 -5.09 -7.63 0.00 -7.63 0.00 -5.09 7.63 -30.52 AR10 -15.19 -18.98 -7.59 -7.59 -3.80 0.00 0.00 -3.80 0.00 -7.59 0.00 -7.59 -11.39 -60.74 Mean -31.38 -28.94 -27.37 -18.77 -18.25 -14.00 -15.23 -13.08 -11.39 -14.69 -4.93 -12.43 -20.84 -69.61 Median -30.25 -27.95 -21.01 -9.22 -15.03 -11.93 -13.92 -6.89 -8.11 -9.97 -2.70 -9.21 -29.92 -73.44 Std. Dev 17.84 32.14 30.30 21.81 23.26 20.55 19.72 23.99 23.13 29.27 30.08 34.16 28.87 46.00

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100 Appendix E (Continued) Table E-4. Oxygen Production at experimental light intensities Amphistegina radiata (nmoles hrP P-1 P PmmP P-2P P) Light Intensity (umol photon mP P-2P PsP P-1P P) Specimen 0 0.96 3.13 10.91 17.45 36.5 48.4 92.1 174.9 233 542 779 1288 0 AR01 -2.79 2.23 2.23 -0.56 2.23 0.56 1.12 1.12 1.12 3.91 1.12 1.68 -3.35 -5.58 AR02 -3.97 -4.63 -1.98 -0.66 -1.32 0.00 -1.32 0.00 0.66 0.00 2.65 1.98 -3.31 -10.58 AR03 -1.87 -3.75 -2.50 -1.87 -1.87 -1.87 -1.87 -0.62 -1.87 -1.25 -0.62 -1.25 -2.50 -8.74 AR04 -1.33 -1.33 -0.66 1.33 0.00 1.99 0.66 3.32 3.32 1.99 5.97 5.97 6.63 0.00 AR05 -3.51 -8.43 -6.32 -5.62 -4.92 -4.92 -4.92 -4.92 -4.21 -5.62 -3.51 -4.92 -3.51 -11.94 AR06 -9.92 0.66 -11.24 -4.63 -7.28 -4.63 -3.31 -5.95 -5.95 -7.28 -9.26 -9.92 -7.28 -18.52 AR07 -5.74 -7.47 -5.74 -6.32 -4.60 -4.02 -5.17 -4.60 -4.02 -4.60 -1.15 -5.17 -5.17 -8.62 AR08 -4.25 -4.86 -3.64 -1.22 -2.43 -1.82 -1.82 -1.82 -1.82 -2.43 -0.61 -1.22 -4.25 -10.93 AR09 -3.55 -3.11 -2.67 -1.33 -1.33 -1.33 -0.89 -1.33 0.00 -1.33 0.00 -0.89 1.33 -5.33 AR10 -2.43 -3.03 -1.21 -1.21 -0.61 0.00 0.00 -0.61 0.00 -1.21 0.00 -1.21 -1.82 -9.70 Mean -3.94 -3.37 -3.37 -2.21 -2.21 -1.60 -1.75 -1.54 -1.28 -1.78 -0.54 -1.49 -2.32 -8.99 Median -3.53 -3.43 -2.58 -1.27 -1.60 -1.58 -1.57 -0.98 -0.91 -1.29 -0.30 -1.21 -3.33 -9.22 Std. Dev 2.45 3.31 3.70 2.47 2.74 2.33 2.17 2.89 2.84 3.40 3.96 4.41 3.85 4.86

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101 Appendix E (Continued) Table E-5. Metabolic Scope and factorial scope for Amphistegina radiata Specimen Normalized to Chl a Norma lized to surface area Factorial scope AR01 25.7 2.8 2.0 AR02 58.0 6.6 2.7 AR03 67.7 6.9 4.7 AR04 -9.3 -1.3 0.0 AR05 92.1 8.4 3.4 AR06 61.4 8.6 1.9 AR07 21.4 2.9 1.5 AR08 59.5 6.7 2.6 AR09 10.2 1.8 1.5 AR10 45.6 7.3 4.0 Mean 43.2 5.1 2.4 Median 51.8 6.7 2.3 Std. Dev 30.6 3.3 1.4 Table E-6. Derived parameters from Photosynthesis/Irradiance curves Normalized to Chl a Normalized to surface area Specimen PB BmaxB B AlphaIB BkB B PB BmaxB B AlphaIB BkB B AR01 AR02 34.95 3.16 11.05 3.98 0.37 10.91 AR03 4.31 0.07 61.74 AR04 49.18 0.27 179.237.10 0.04 190.55 AR05 AR06 AR07 12.50 0.21 60.83 1.68 0.03 61.09 AR08 20.51 2.74 7.49 2.30 0.32 7.26 AR09 17.13 1.27 13.53 2.99 0.22 13.50 AR10 11.33 1.29 8.78 1.81 0.21 8.84 Mean 21.42 1.29 48.95 3.31 0.20 48.69 Median 17.13 1.27 13.53 2.65 0.21 12.20 Std. Dev. 15.53 1.25 62.32 2.04 0.14 72.45


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Walker, Robert A.,
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Photosynthesis and respiration in five species of benthic foraminifera that host algal symbionts
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by Robert A. Walker.
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[Tampa, Fla.] :
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2004.
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Thesis (M.S.)--University of South Florida, 2004.
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ABSTRACT: Oxygen production and consumption were measured in five species of benthic foraminifers using a "Clark-type" oxygen electrode. Net photosynthesis and respiration were calculated and normalized to both μg Chl a and mm upper surface area for the chlorophyte-bearing soritid foraminifers, Archaias angulatus and Cyclorbiculina compressa, and the diatom-bearing amphisteginids, Amphistegina gibbosa, A. lessonii and A. radiata. Photosynthesis/Irradiance curves were generated by fitting data to the hyperbolic tangent equation P = Pmax tanh (α I/ Pmax). Derived photosynthetic parameters, Pmax, α, Ik were found to correspond to the general responses of the endosymbiont taxa. Chlorophyll concentration was found to be significantly lower in Cyclorbiculina compressa than in the other four species. Maximum O production (Pmax) when normalized to Chl a was 3-4 times higher in soritid species than in amphisteginids. Photosynthetic efficiency (α) was significantly higher in Amphistegina gibbosa and A. lessonii than in the soritids. Mean Ik, which indicates approaching light saturation, was 13 and 26 μmol photon m 2sec1 respectively for A. gibbosa and A. lessonii compared with 95 and 119 μmol photon m2sec1 respectively for Archaias and Cyclorbiculina. Calculated P/I data were to variable for Amphistegina radiata to estimate reliable α and Ik values. Factorial metabolic scope, which indicates potential for activity was only 2-6 for amphisteginids versus 9-16 for soritids. Annual primary production was estimated to be 285 mmoles O m2 of habitat for A. angulatus, 9.3 mmoles O m2 of habitat for C. compressa and 15.3 mmoles O m2 of habitat for Amphistegina lessonii. Pmax values for Amphistegina gibbosa fluctuated at the compensation point and did not indicate significant oxygen production. Pmax values for Amphistegina radiata failed to reach the compensation point and net oxygen production was not recorded.
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Adviser: Pamela Hallock Muller.
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irradiance.
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