The Global Ecology of C 4 Grasses. Colin Osborne University of Sheffield
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1 The Global Ecology of C 4 Grasses Colin Osborne University of Sheffield
2 The C 4 syndrome changes environmental responses of photosynthesis Björkman (1971) Photosynthesis & Photorespira/on (eds. Hatch, Osmond, Slatyer)
3 But ecological adapta<on also has large effects... Pearcy & Ehleringer (1983) Plant Cell Environ
4 Evolu<onary perspec<ve changes interpreta<on of global pa?erns Edwards & Smith (2010) PNAS
5 C 4 grass lineages differ hugely in their biology Trait value Fig. 5 Comparisons of main traits between Chloridoideae and Panicoideae. Hui Liu Liu et al. (2012) Oecologia
6 Unraveling effects of ecology & evolu<onary history
7 How does C 4 photosynthesis interact with other determinants of plant growth?
8 Growth rate is a fundamental axis of varia<on in plant ecology compesson ferslity gradients trait correlasons e.g. N content Sssue turnover growth growth growth trade- offs survival defence storage
9 How does C 4 photosynthesis interact with other determinants of plant growth? ul/mate physiological innovason growth ecology evolusonary history
10 How does C 4 photosynthesis interact with other determinants of plant growth? proximate ul/mate photosynthesis physiological innovason allometry growth ecology development evolusonary history
11 Differen<al growth is major driver of community structure in global dynamic vegeta<on models DGVM C 4 grass PFT Woodward et al. (2004) Phil. Trans. R. Soc. B
12 South Africa Grass lineages occupy different fire environments Andropogoneae lineage Mesic savannas High fire frequency ArisSdoideae lineage Hot, arid grasslands Heavy grazing Vernon Visser Visser et al. (2011) J. Biogeography
13 Do members of different grass lineages regrow at different rates aier fire? Andropogoneae C 4 Brad Ripley Paniceae C 3 ArisSdoideae C 4 Danthonioideae C 3 Tarryn MarSn
14 Do members of different grass lineages regrow at different rates aier fire?
15 Regrowth rate varies among grass lineages and is an adapta<on to fire Ln rate of recovery (mg d -1 ) a C 4 C 3 P < 0.05 Number of fires Rate or recovery Panicum aequinerve Alloteropsis eckloniana Panicum ecklonii Heteropogon contortus Themeda triandra Hyparrhenia hirta Pentaschistis pallida Pentaschistis curvifolia Merxmuellera disticha Karroochloa curva Aristida diffusa Arsitida junciformis Ln number of fires P- A ChrisSn Vernon Visser
16 Ecological adapta<on and shared evolu<onary history have major effects on regrowth aier fire. Larger sample of species needed to dis<nguish hypothesized C 4 effect from these other effects.
17 Larger sample of species needed to dis<nguish hypothesized C 4 effect from these other effects. Sam Taylor Taylor et al. (2009) New Phytol
18 Are there general effects of C 4 photosynthesis on growth rate among grasses? Chris Bennec Emily Mockford Rebecca Atkinson Poster P39
19 Are there general effects of C 4 photosynthesis on growth rate among grasses? C 3 /C 4 photosynthesis 5 EvoluSonary history Climate (MAT/MAP) Habitat (Open/Wet) Life history ~450 spp = 4,500 plants Growth form DomesScaSon status
20
21 Rela<ve growth rate in >440 grass species fixed species um monococcum var. monococcum Vulpia myuros Stenotaphrum secundatum Sacciolepis striata Arthraxon prionodes Leptochloa virgata Oryza grandiglumis Trichoneura grandiglumis Sporobolus pyramidalis Aristida adscensionis Anthephora pubescens Tricholaena monachne Eragrostis airoides Setaria parviflora Melinis ambigua Spartina pectinata Digitaria seriata Brachiaria Hordeum subquadripara vulgare subsp. Secale vulgare vavilovii Diheteropogon hagerupii Cynodon barberiavena fatua Buchloe dactyloides Ehrharta calycina Bouteloua curtipendula Lolium multiflorum Bothriochloa pertusa Panicum laetum Eragrostis papposa Muhlenbergia racemosa Tribolium echinatum Bothriochloa saccharoides Stipa viridula Eriochloa meyeriana Bothriochloa insculpta Paspalum conspersum Lolium Dichanthelium perenneacuminatum Alopecurus subsp. spretum myosuroides Eragrostis patula Dactylis glomerata Calamovilfa longifolia Setaria leucopila Chloris radiata Digitaria didactyla Rostraria cristata Homopholis Microstegium proluta vimineum Triticum timopheevii Sorghum angustum Panicum aequinerve Panicum phragmitoides Muhlenbergia rigens log.m Pennisetum sieberianum Eragrostis pectinacea Eragrostis superba Digitaria monodactyla Echinochloa crus galli Dactyloctenium Sorghum radulans bicolor subsp. Eleusine verticilliflorum coracana Dactyloctenium Sorghum giganteumarundinaceum Pennisetum glaucum Brachiaria lata Coix lacryma jobi Digitaria ciliaris Chionachne gigantea Panicum miliaceum Sorghum amplum Hordeum Panicum spontaneum miliaceum var. Coelachyrum ruderalepanicum piercei dichotomiflorum Eragrostis pilosa Cenchrus brownii Setaria italica Panicum capillare Digitaria setigera Cenchrus echinatus Echinochloa picta Enteropogon Pogonarthria prieurii squarrosa Panicum sumatrense Danthoniopsis Paspalum dinteri malacophyllum Sorghum Zea bicolor mays subsp. parviglumis Digitaria longiflora Echinochloa esculenta Cenchrus setigerus Eragrostis tef Zea mayssorghum ecarinatum Paspalum hartwegianum Leptochloa fusca Eragrostis cilianensis Paspalidium desertorum Fingerhuthia africana Eragrostis intermedia Stipa capensis Digitaria abyssinica Setaria verticillata Digitaria sanguinalis Ischaemum Dactyloctenium rugosum aegyptium A. speltoides ligustica Melinis repens Sorghum timorense Urochloa mosambicensis Secale cereale Digitaria fuscescens Panicum stapfianum Tragus racemosus Cenchrus incertus Avena sativa Panicum antidotale Sorghum propinquum Pennisetum violaceum Digitaria bicornis Lintonia nutans Paspalum vaginatum Eragrostis rufescens Oryza glaberrima Sporobolus fimbriatus Apluda mutica Oryza barthii Echinochloa crus pavonis Eragrostis eriopoda Chloris elata Cynodon transvaalensis Heteropogon contortus Bouteloua repens Eleusine indicatrichloris crinita Eragrostis mexicana Zoysia japonica Eragrostis unioloides Stipagrostis uniplumis Paspalum distichum Setaria faberi Eriachne Panicum aristidea pseudowoeltzkowii Cenchrus pilosus Enneapogon scoparius Bouteloua gracilis Triticum Stipagrostis aestivumhirtigluma var. hirtiglum Whiteochloa capillipes Digitaria brownii Polypogon fugax Echinochloa stagnina Oxychloris scariosa Iseilema macratherum Hordeum comosum Leptochloa panicoides Dinebra retroflexa Eragrostis capensis Eleusine kigeziensis Stipagrostis Pennisetum ciliata clandestinum Enteropogon sechellensis Panicum bisulcatum Digitaria debilissetaria viridis Bothriochloa Stipagrostis barbinodis hochstetteriana Paspalum dilatatum Cynodon dactylon Pennisetum setaceum Dasypyrum villosum Chloris gayana Echinochloa Vaseyochloa muricata multinervosa Panicum hirticaule Leptochloa dubia Cymbopogon citratus Poa alpina Aristida diffusaaegilops crassa Leymus triticoides Digitaria milanjiana Panicum Cymbopogon virgatum jwarancusa Phalaris subsp. jwarancusa arundinacea Leymus karelinii Muhlenbergia porteri Sporobolus helvolus Tripogon jacquemontii Pennisetum lanatum Setaria barbata Digitaria ternata Polypogon Stipagrostis chilensishirtigluma Sporobolus var. patulaindicus Digitaria insularis Tetrachne dregei Triraphis schinzii Aeluropus macrostachyus Poa annua Digitaria exilis Eremopyrum triticeum Arundinella hirta Festuca Triticum orthophylla monococcum subsp. Elymus aegilopoides tibeticus Bothriochloa springfieldii Paspalum floridanum Eragrostis variabilis Paspalum scrobiculatum Aristida funiculata Triticum baeoticum Dichanthelium scoparium Eulalia aurea Loudetiopsis kerstingii Oplismenus hirtellus Sporobolus fertilis Nassella pubiflora Dichanthium aristatum Bothriochloa bladhii Alloteropsis cimicina Briza maxima Panicum trichanthum Schizachyrium sanguineum Poa palustris Erioneuron pulchellum Loudetia phragmitoides Vulpia microstachys Eragrostis porosa Aristida ternipes Iseilema membranaceum Aristida barbicollis Elymus mutabilis Panicum lanipes Elymus dentatus Brachypodium Heteranthelium pinnatum Acroceras piliferum zizanioides Tristachya superba Loudetia Achnatherum simplex calamagrostis Piptochaetium napostaense Panicum laxum Muhlenbergia Leersia rigida drepanothrix Panicum natalense Paspalum Calamagrostis simplex canadensis Calamovilfa gigantea Elymus glaucissimus Desmazeria sicula Panicum tenellum Paspalum unispicatum Oryza rufipogon Cymbopogon commutatus Triticum dococcoides Bromus sterilis Piptochaetium avenaceum Melica ciliata Pennisetum alopecuroides Eriochloa sericea Psathyrostachys Phalaris junceacoerulescens Oryzopsis miliacea Poa sieberiana Pennisetum purpureum Paspalum tenellum Panicum subalbidum Andropogon Chasmanthium hallii Rytidosperma latifolium unarede Lasiacis sorghoidea Arundinella hookeri Poa arachnifera Lygeum spartum Zuloagaea bulbosa Paspalum fimbriatum Tripogon filiformis Panicum amarum Spodiopogon Arthraxon sibiricus lanceolatus var. Panicum echinatus rigidulum Halopyrum mucronatum Ehrharta Brachypodium erecta phoenicoides Oryza meyeriana Sacciolepis chevalieri Stipagrostis obtusa Poa cita Oryza latifolia Panicum dregeanum Danthonia spicata Panicum pygmaeum Festuca arundinacea Panicum hallii Koeleria cristatastipa tenuifolia Eleusine tristachya Paspalum haumanii Orthoclada laxa Elionurus euchaetus Hymenachne amplexicaulis Astrebla pectinata Poa compressa Tribolium hispidum Oryza punctata Puccinellia distans Oryza eichingeri A. semialata Semialata Odyssea mucronata 2 Hordeum marinum Melica altissima Panicum pansum Aristida divaricata Gastridium ventricosum Cortaderia selloana Brachypodium distachyon Leersia oryzoides Stipagrostis plumosa Stipa tenuissima Paspalum notatum Andropogon gerardii Bromus inermis Ischaemum Taeniatherum afrum caput medusae Lasiacis divaricata Echinochloa pyramidalis Stipa parviflora Aristida behriana Digitaria gayana Elymus pungens Dichanthelium acuminatum Dichanthium foveolatum Alopecurus textilis Sorghastrum Muhlenbergia nutans A. glomerata semialata Eckloniana Phleum pratense Panicum nervatum Elymus farctus Agropyron fragilestipa barbata Axonopus compressus Dichanthelium Aristida glaziovii oligosanthes Imperata var. oligosanthes cylindrica Poa maniototo Merxmuellera Paspalum stricta conjugatum Panicum queenslandicum Muhlenbergia capillaris Stipa spartea Sporobolus wrightii Stipa brandisii Calamagrostis arundinacea Eragrostis Schedonnardus lugens Dichanthelium paniculatus scabriusculum Chasmanthium laxum Panicum Panicum bergii rigidulum subsp. Aristida rigidulum congesta Panicum millioides Panicum turgidum kmannia syziga Triplachne nitenstipa sareptananisetum basedecurus magella Danthonia parryi Aristida hordeacea Melica macra Setaria megaphylla Danthonia californica Eragrostis gummiflua Brachypodium sylvaticum Paspalum minus Poa iberica Miscanthus oligostachyus Poa gilgianafestuca lenensis Paspalum pumilum Cymbopogon pospischilii Paspalum pubiflorum Arrhenatherum Arctagrostis album latifolia Stipagrostis Anthoxanthum zeyheri odoratum Poa aridafestuca spectabilis Elymus patagonicus Anthenantia rufabriza uniolae Stipa roylei grostis stolonifeaxonopus sppbromus mangofestuca kingii Panicum pilosum Glyceria grandiscciolepis africaanthonia sericeuropus lagopoi nthonia decumb Briza subaristat Bromus erectus Poa arctica nthonia compresperma semian Festuca roemer Festuca callieri spalum thunbeoleataenia ance orobolus gigantdosperma pilospoa colensoi icum clandestin oceras amplect ogon monspelinicum brevifoliuaira cupaniananna arundinaceustachys petraertococcum patefestuca hallerifestuca petraeaymus canadenshionochloa rubdichelachne rar Aristida adoensimarthria altissithelium depaupnardus strictaestuca californic ciolepis cymbiamicrochloa caffrsleria heufleriatriodia pungensachather paradpanicum prioniti ysanolaena latifdolepis mitchelkoeleria nitidulahanthelium leiberidolepis xerop
22 Are there general effects of C 4 photosynthesis on rela<ve growth rate among grasses? effect size significance C 4 photosynthesis +++ *** Annual life history +++ *** DomesScaSon ++ * Sod- forming growth + ** Open habitat preference + * Higher precipitason * Shared evolusonary history ++ ***
23 relasve growth rate = specific leaf area x leaf mass raso x net assimilason rate RGR = SLA x LMR x NAR Leaf area per unit leaf mass Leaf mass per unit plant mass Growth per unit leaf area
24 relasve growth rate = specific leaf area x leaf mass raso x net assimilason rate RGR = SLA x LMR x NAR C 4 photosynthesis +++ Annual life history DomesScaSon ++ Sod- forming growth ++ Open habitat preference PrecipitaSon
25 Why is specific leaf area higher in C 4 than C 3? SLA = 1 leaf density * leaf thickness EvoluSon of C 4 photosynthesis led to thinner leaves in grasses P- A ChrisSn Erika Edwards thickness C4 OS thickness C4 IS C 3 C 4 C 3 C ChrisSn et al. (2013) PNAS
26 The greater efficiency of C 4 leaves is linked to changes in the alloca<on of growth Sage & Pearcy (1987) Plant Physiology Long (1999) in C 4 Plant Biology (eds. Sage & Monson) Thinner leaves Greater leaf area Faster growth Greater root mass allocason Access to soil water/n?
27 Why doesn t C 4 photosynthesis boost the net assimila<on rate? Similar whole- plant net assimilason rate per unit leaf area Faster photosynthesis Greater root respirason? Faster leaf turnover?
28 The effect of C 4 photosynthesis on growth is mediated via changes to leaf thickness and root alloca<on
29 These changes in alloca<on exert major effects on growth, reminding us that photosynthesis acts within the context of a whole organism
30 Acknowledgements Rebecca Atkinson Emily Mockford Chris Bennec Mark Rees Rob Freckleton Ken Thompson Pascal- AnSone ChrisSn Erika Edwards Brad Ripley Tarryn MarSn Sally Archibald Vernon Visser Sam Taylor Hui Liu David Simpson Derek Clayton Knowledge Now Limited
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