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Production Ecology and Ecophysiology of Turf Algal

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Production Ecology and Ecophysiology of Turf Algal YOÀ io'l Production ecology and ecophysiology of turf algal communities on a temPerate reef (West Island, South Australia) Margareth CoPertino Department of Environmental Biology The UniversitY of Adelaide Thesis submitted for the degree of Doctor of PhilosoPhY l|llarch2002 Table of Contents V LIST OF FIGURES... x LIST OF TABLES..... XIII LIST OF PLATES..... xIv LIST OF ABBREVIATIONS XVI ABSTRACT XVIII xIx ACKNOWLEDGEMENTS ............... I 2 2 4 CHAPTER I INTRODUCTION 1.1TuRFALGALcoMMIJNITIES.'...'... """"""""""'4 """"""""7 1.2 PRIMARY PRoDUCTIoN l0 1.3 BouNDARYLAYERANDDENSITYEFFEcrs """"""""""""" 12 1.4 THE BASIS FOR PRODUCTION: PHOTOSYNTHESIS""""""" """"""""""""' 14 1.5 PHoToAccLMATIoN AND PHoroINHlBlrIoN """"""""""" 19 1.6 CoNCLUSIONS......'.....' """"""" CHAPTER II PRODUCTIVITY RATES AI\D BIOMASS ........""' """20 20 2.1 INTRODUCTION 24 24 2.2.1 Site description 2.2.2 Experimental design and settlement plates 2.2.3 Biomass 2.2.4 CommunitY structure 2. 2. 5 Primary productivity...'......... ;... " "' 2.2.6 Statistical analYsis 2.3.1 Light and temperature ..'.....-'..... 2. 3. 2 CommunitY structure 2. 3. 3 Biomass standing crop...........'.. 2. 3.4 ProductivitY rates.. " 2.4 DßcussloN................ 2.4.1 SeasonalitY and dePth 2.4.2 Importance of turfs on temperqte reefs """"""' 2.4.3 Aòomparison lo tuds on tropical reefs """""" 2.5 CoNcLUSIoNS .....'.'..... 72 CHAPTERIIICIIAI\GESINPHOTOSYNTHETICPARAMETERS................ 72 3. I INrnouucrloN........... 78 3.2 R¡VTEW ON P-E CURVE AND BIOPHYSICAL PARAMETERS '.....'...'.... 84 84 86 3. 3.2 Statiscal analYsis 11 .. 88 88 3.4. I Light and temperature .......'..'.... 88 3.4.2 Atl data sets: dffirences 'tetvveen seasons and depths """ 89 3.4. 3 Biomass effects........ 94 3.4.4 Morning and afternoon curves .'..'... 98 3.4.5 Morning curves for biomass classes 98 Differences between biomass classes ........... 99 Differences between depths....'.'..'.. 99 Differences between seasons t02 3.5 DßCUSSION 102 3. 5. 1 Depth effects ....... t0s 3. 5. 2 Seasonql effects.........'. -. - t09 3. 5. 3 Biomass effects........ 111 3.5.4 P-E curve hysteresis effects 115 3.5 CoNct-usloNs r18 CHAPTER IV VARIABILITY IN PHOTOSYNTHETIC EF'FICIENCY................" 118 4. I INTRoDUcrloN........... 122 4.2 A nevmw oN CHL,4 FLUoRESCENCE PARAMETERS."""""""" 124 4.3 Msrsoos ................... 124 4.3.1 Daily cycle of Yield in situ. 124 125 Turfs on natural rock substratum..... t26 4.j.2 Daily cycle of yield in outdoor tanks ..'......... 126 Experiment l: Plates on different days ...'......'.... t27 Experiment 2: Plates vs boulders, 2m vs 4m, high light vs low light" 127 4.3. 3 PAM fluorometry ... 128 4. 3.4 Yield-líght cunes..........'. 130 4.3.5 Statistical analYsis r32 132 4.4.1 Dffirences betvveen seasons, depths and plate age"""""""' t37 4.4.2 Dffirences betvveen tud comPonents 139 4.4.3 Dffirences b etvv een morning and afternoon curves.........'.. 144 4.4.4 Daily cycle ofyield in outdoor tanks 145 Plates on different days..'..'.......... t47 Plates versus boulders, 2m versus 4m, high light versus low light ' 151 4.5 DtscussIoN................ I5I 4.5.1 Daily cycle of effective quantumyield."""""' 153 4.5.2 Daily cycle under very low PPFD 156 4. 5. 3 Differences b etw een tud components .........'.. " " 160 4.5.4 Depth effects .......... 162 4. 5. 5 Seasonal effects......... 166 4.5.6 The high variability around curves........-.. 167 4.6 CoNCLUSIoNS ............. CHAPTERV PHOTOINHIBITION AND PIIOTOACCLIMATION........... """"" 170 170 t75 5.2 METHODS 175 5. 2. I Transplants ............'... 175 Transplantation . Transplantation ubshatum) 176 178 5.2.2 Changes in Yield (Y') 179 5.2.3 Changes in Photosynthetic parameters r80 5.2.4 Statistical analysis 181 5.3 RESULTS. 181 5. 3. 1 Transplant ation of settl ement p lates ....'...... 181 Changes in effective quantum yield in the first 48-hours.. 183 Changes in effective quantum yield over 15 days.. 191 Changes in photosynthetic parameters after 15 days """""""' rs (turf components) """"' 193 5.3.2 Transp 193 eld in the first 48-hours" """"' Changes in 199 Changes in eld over 7 days""""""' """""' 205 5.4.1 Photoinhibition/ relaxation period ' """ """" 205 Turf communities on settlement plates """' 210 5.4.2 Photoacclimation period.....'.."""' """" """'211 Tìme course: changes in effective quantum yield"""""""' Changes in photosynthetic parameters' """""""'215 CHAPTERVI SYNTIIESIS ""'2I9 220 6.1 PRODUCTTVITY RATES 222 6.2 THE PHOTOACCLIMATION MECHANISM.' 227 PHOTOACCLIMATION' PRODUCTION AND GROÌúTH 6.3 A BALANCE BETWEEN 23t 6.4 CoNcLUSIoNS AND FUTURE DIREcrloNS.' 235 259 260 262 265 273 274 lv List of Figures Figure 2.1 Location of West Island in Encounter Bay, Victor Harbour, 800 metres off shore. The insert shows the island's position in relation to Australia.. """"' 28 m (blue Figure 2.2 Average maximum PPFD per month (right axis) at the depths of 4 m (red line) and l0 line), temperature per month (left axis, black line) and day length in hours (numbers at the top) 40 during the studied ieriod' Complete table of values in Appendix 4""""""""" """"' during Figure 2.3 Meat(+SD) percentage cover of turf algal components present on plates at 4 and l0 m, (pink)' the period of Sepìember lsgg and June 2000. Blue green algae (blue), encrusting coralline brown filamentous red ired), filamentous brown (brown), filamentous green (green) and encrusting Figure 2.4 Mean (+SD) percentage cover of turf algal components present on plates at 4 and 10 m, collected during'tvtarch 2000, from age 3 to 18 months. Blue green algae (blue), encrusting (green) coralline (pink), hlamentous red (red), filamentous brown (brown), hlamentous green and 45 encrusting brown (black).. """' Figure 2.5 Mean (+SD) number of juvenile macroalgae (from each component) present in turf algal communities on settlement plates at 4 and lOm, during the period of Septembet 1999 and June sheet like 2000. Articulated coralline (pink), brown foliose or branched (brown), green branched or (green), red foliose or branched (red)'.......'. .. """""""""' 46 Figure 2.6 Mean (+SD) number of juvenile macroalgae (for each component) present in turf algal communities on settlement plaies at 4 and 10m (during March 2000) from age 3 to 18 months' Articulated coralline (pi"k); brown foliose or branched (brown), green branched or sheet like (green), red foliose or bìanched (red)...'.'....... """""""""' 47 (blue) Figure 2.7 Meanbiomass (+SD) of turfs growing on settlement at the depths of 4 m (red) and 10 m during the studied period. (a) Biomass per month and depth; (b) biomass per plate age and depth, recorãed during March 2000............ """""" 54 for all Figure 2.8 Biomass specihc productivity as a function of standing biomass, for 4-m and l0-m turfs, months. Green dots, spring; ,"á dotr, surnmer, brown dots autumn. Relationship approaches an exponential decline......... """"' 55 Figure 2.9 Difference between seasons and depths (Tukey p < 0.05) for biomass specihc net productivity (a) and productivity on an areal basis (b). vlean (y-axis), variance (bubble area) and groups of 56 À"un. 1i-u*is) in accord to the test. """""" Figure 3.1 Model of light-saturation for net photosynthesis, measured as Oz evolved or C fixed, versus ppFD (p-E curvá). Pm, maximum photosynthesis; Pg, gross photosynthesis; Pn, net photosynthesis; Rd, daik respiraíion; Éc, üght compensation poinq Ek, light saturation index; a, maximum light utilization coefficient.... """""'82 the Figure 3.2 Models of adjustment of the photosynthetic unit to extreme low and high light by changing light. number or size oireaction centres. (a) Fixed antenna size, increasing number of traps in low (b) Fixed number of traps, changing antennr size. (c) Increasing both number and antenna size' Èígh üght acclimated in red, low iight acclimated in blue. Modihed from Ramus 1981. .'.........'.'. 83 a 20 Figure 3.3 Typical p-E curves measured at the depth of 4 m with exponential fitting obtained from >0.8 90 hour áðployment using 4 m (red) and 10 m turfs (blue). Only curves with 12 were used...'.'.. Figure 3.4 photosynthesisJight relationships for each season estimated from an exponential model, for turfs growing at deptñs of (a) 4 m and (b) l0 m measured at 3-4 m. spring (green), summer (red), autumn (black) and winter (blue). """""""" 9l growing at Figure 3.5 Effect of biomass on Pm values measured at different seasons and days, for turfs summer depths of (a) 4 m and (b) 10 m. Different coloured dots represent seasons: spring (green)' (tà), u.ttotntt (black) aná winter (blue)' '.....'.'. """"""""'92 deployment site Figure 3.6 Example of hysteresis in P-E curves on a day of high maximum PPFD at the (4m), with aft"rnoon decrease in photosynthetic response of 4-m (red) and lO-m (blue) turfs' Morning curyes (full line and dots) htted for data collected between 01:00 and 12:59 h, and afternoon curves (àashed line and open dots) for data collected after l3:00h.. """""'96 for turfs Figure 3.? photosynthesis-light relationships for each season estimated from exponential model, (g¡een), growing at 4 m(a, bfand l0 m depths (c, d), with high and low standing biomass. Spring ,n-*"i (red), autumn (black), winier (blue). .'."""' """' 100 Settlement Figure 4.1 Schema o during December 1999' (red) plates þ) and of 2 m (green) and 4 m to an tl ' """' 131 "*por"å'1t growing on plates at Figure 4.2qffecfive quantum yield as a function of PPFD (Y-E curves) for turfs for the depths of 4 mlred) u.tá tO m (blue) measured from June 1998 to June 2000. Parameters 134 curyes are in Table 4.1. ....'.'....." """""""" old Figure 4.3 yield-light curves for turfs growing on 3-month (full line and filled dots) and l2-month (a) plates (dashãd üne and open dots) at dãpths of4 m (red) and 10 m (blue) measured in spring '9S 136 and (b) autumn 99' Paiameters of the curyes in Table 4'3"""""""' """""""""" of 2 m (green) Figure 4.4 yield-light curves for different components of natural turf communities at depths 138 and 4 m (red) measured during March 2000 growing on plates at Figure 4.5 1'-E curves based on morning (blue) and afternoon data (red) for turfs curves are in depths of 4 m (left) and 10 m (righÐ from June 1998 to May 1999.
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