Posted on Authorea 24 Jun 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159301657.74157566 — This a preprint and has not been peer reviewed. Data may be preliminary. n h eaiepootosaogteefatos h loaino ofla rcin slkl related stress likely and is reproduction, fractions growth, foliar to to related P functionally of fractions are P allocation fractions foliar The the these of fractions. because concentrations strategy, these the including small life-history among both cells and to in proportions leaf differences phospholipids relative reflect in acids, the will fractions nucleic and conditions environmental P (i.e. to major due compounds several or organic 2005). of P-containing Bloom, concentrations (Veneklaas various & esters the phosphate (Epstein and of concentration (Pi) sum P P foliar the inorganic low is a concentration having P by availability Foliar P soil low (Hou (Kooyman to ecosystems Australia respond terrestrial in generally natural widespread is in availability pervasive P is production plant Thomas above-ground 1997, (Lambers Conroy, ecology (Conroy & physiological world the plant phos- throughout in to forests topic adapted plants important in an mechanisms Veneklaas is underlying soils their (P)-impoverished and allocation phorus nutrient of strategies Understanding P soil to patterns response INTRODUCTION nutrient-allocation in the strategies that different conclude revealed We attenuata strategy. similar. growth B. were more contrasting slower-growing allocated fractions their and sessilis other matched B. sessilis which Faster-growing but P availability, B. acid did, fractions. nucleic N opportunistic attenuata foliar and B. faster-growing P both decreased than in other with species B. with acids correlated both correlation of nucleic positively in no was concentration concentrations to was N species N P there and both but total P of concentrations, total foliar rate N foliar The growth total The relative and fractions. substrates. foliar The all availability. P-containing similar on P had to attenuata decreasing availability. species B. allocation with two of P P on that The different of than occurs with fractions. greater patterns and substrates biochemical was distinct fire sessilis on major by plants but to killed with concentrations, allocated resprouts is experiments P P which which pot and total (Proteaceae), sessilis, out concentrations, B. nutrient attenuata carried We foliar opportunistic Banksia measured faster-growing limestone. slow-growing We with or in sand, laterite (N) over deep nitrogen sand on shallow occurs and naturally (P) and phosphorus fire of after use the compared We Abstract 2020 24, June 4 3 2 1 Lambers Han Zhongming to adaptation and habitats rate low-phosphorus growth in differing and sessilis attenuata Banksia Banksia in patterns nutrient-allocation Foliar nvriyo etr Australia Western of Australia University Western of University The University Agricultural Jiangxi University Agricultural Jilin 3 tal. et 02.Popou-moeihdsislmttegot n il fcos atrsand pastures crops, of yield and growth the limit soils Phosphorus-impoverished 2012). , 1 ini Shi Jianmin , tal. et tal. et 06.Mroe,arcn eaaayi hwdta iiainof limitation P that showed meta-analysis recent a Moreover, 2006). , 02.Teeoe aito nfla ocnrto mn ln species plant among concentration P foliar in variation Therefore, 2012). , 2 iynPang Jiayin , tal. et 90 Fujita 1990, , 3 tal. et 1 iYan Li , tal. et 06 icraRse u,21) n plants and 2016), Bui, & Rossel Viscarra 2016, , 4 arc Finnegan Patrick , 03 ebr ons 95 Seneweera 1995, Fownes, & Herbert 2003, , tal. et 4 n Hans and , 00.Lwsoil Low 2020). , tal. et 2006, , Posted on Authorea 24 Jun 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159301657.74157566 — This a preprint and has not been peer reviewed. Data may be preliminary. oeo hs rtaeeehbtavr ihpooytei -s ffiiny(PE Denton (PPUE, efficiency P-use photosynthetic high very a exhibit Lambers Proteaceae these fast of relatively (Brooks some exhibit area 2005). leaf (Denton Australia unit concentrations Lambers, south-western per P photosynthesis & from leaf low (Shane species extremely Proteaceae particles having Lambers despite of soil photosynthesis, leaves from area-based mature of P rates that desorb striking to is carboxylates It possess low-molecular-weight likely on vegetation of they the thus, amounts of status; (Hayes component Australia pool, P important south-western an RNA al. soil in are the low soils Proteaceae Within long-term P-impoverished Non-mycorrhizal severely experienced 2002). limitation. have Roche, P landscapes to La adaptations ancient & in (Geider species proportion Tree largest fraction. the P (Reef largest far 4:1 the Elser by of is 2007, stoichiometry RNA rRNA Hamilton, N:P with synthesis P, & an protein have (Elser organic the allocate rates acids of meet must growth to Nucleic organisms (rRNA) rapid rates because 2002). RNA the growth N, Elser, ribosomal support fast P-rich for to that to than needed P proposes P (Reef of demands which for ratios proportion requirement (GRH), N:P greater stoi- greater Hypothesis disproportionately low ecological proportionally Rate a have a of Growth rates with field the growth associated by rapidly-emerging are fast explained the with been and has species investment, pattern that P shown and have rate genus chiometry growth same between P the relationship in allocate species The plants two climate How in Mediterranean a P with use adaptive. distribution ecosystems efficiently unclear. P-impoverished likely their to extremely remains are in strategies with in strategies adaptive differences associated life-history exhibit P contrasting the was available and with fractions that pattern of P gradient concluded allocation foliar strong and P among a chronosequence, their Yan with that the PPUE. chronosequence found along two-million-year high allocated and maintain a be may to Australia, along P P south-western species cellular structural of three proportion P-impoverished to of greater on than a fractions compared growing rather that (PPUE) plants P, suggested efficiency authors P-use that metabolic These photosynthetic found to and soil. (LMA) (2009) P-richer area Kitayama on per the plants & mass with Hidaka in leaf difference a both expected. increased demonstrated no soils be was species that g tropical can there these mg found limitation However, that (1.3–2.1 and P and types. concentration species to forest season, P tree different growing foliar Alaskan between the high four concentrations throughout relatively P of pool foliar fractions largest of prevailing P the proportion the foliar was under investigated P fitness (1983) acid plant nucleic Kedrowski increase to & might acclimate this III to and Chapin fractions, plants 2011). P for Kitayama, foliar mechanism & among (Hidaka Yan important P conditions 2011, of an Kitayama, allocation is & the leaves (Hidaka shift availability in P patterns soil low P-allocation Shifting tolerance. elcmn fpopoiiswt aatlpd n uflpd uigla eeomn (Lambers development resprouter slow-growing leaf The during (Sulpice sulfolipids concentrations and rRNA galactolipids foliar 2012). with low phospholipids by of mainly replacement about brought is habitats tp/flrbs.pww.o.u)adde o rwfs n opeeislf yl ucl Bwn& (Bowen quickly than cycle rate life faster its a complete at and these fast exuding grow and/or not carboxylates does more and http://florabase.dpaw.wa.gov.au/) releasing by (Shi limestone attenuata histories or life laterite different from have but 2005), attenuata Lambers, (Hayes & limestone (Shane roots cluster al. compound produce both 01.Seisi hsfml yial omcutrrosta ffcieymn olPb eesn large releasing by P soil mine effectively that roots cluster form typically family this in Species 2001). , 2020). , tal. et tal. et (Shi hc net oei epros(Shi roots deep in more invests which , aki sessilis Banksia tal. et 02 Sulpice 2012, , 00 Sulpice 2010, , tal. et 00.I otatto contrast In 2020). , 09 ae&Bl,19)adalctsmr ims ocutrrosthan roots cluster to biomass more allocates and 1999) Bell, & Pate 2019, , aki attenuata Banksia sasotlvdolgt edrta cuso hlo adoe aeieor laterite over sand shallow on occurs that seeder obligate short-lived a is tal. et tal. et 04,wielae fPsavdco lnstn ohv lwrtsof rates slow have to tend plants crop P-starved of leaves while 2014), , 04.Ti ihPU nPoeca rmsvrl P-impoverished severely from Proteaceae in PPUE high This 2014). , tal. et .sessilis B. 98 Fredeen 1988, , n h atrgoigseeder faster-growing the and -1 tal. et 2 r as(M) on dpaino h respecies tree the of adaptation no so (DM)), mass dry , tal. et .attenuata B. 00.Ti taeyehne mobilisation P enhances strategy This 2020). , tal. et 09.I togPlmtto cus plants occurs, limitation P strong If 2019). , tal. et 90 Rao 1990, , tal. et srsrce ode ad(FloraBase, sand deep to restricted is 04 Lambers 2014, , tal. et 00,adaeamjrfraction major a are and 2010), , .sessilis B. tal. et tal. et 21)ivsiae oirP foliar investigated (2019) tal. et 99.Consequently, 1989). , 04 n extensive and 2014) , tal. et tal. et (Pate 96 tre & Sterner 1996, , 03 Pate 2013, , 00.This 2010). , tal. et tal. et tal. et tal. et 1991) , 2007, , 2007, , B. B. et et , Posted on Authorea 24 Jun 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159301657.74157566 — This a preprint and has not been peer reviewed. Data may be preliminary. o xeietwscridoti lshuea h nvriyo etr utai,Prh Australia Perth, Australia, Western of University the at glasshouse a in out carried (31 was experiment pot A (30 Australia Western Bay, design Jurien was Experimental near seedling limestone One over growing paper. population, 115 filter coastal S, a on from 22 sown collected on were were pot Australia), experimental Western each into River, transferred King Company, Seed Nindethana both of concentrations 2) P total foliar the availability, decrease. P would soil decreasing With 1) that: hypothesised and We concentrations and in LMA, two fractions concentrations, these P-containing N contrasting in and foliar in availability. P in patterns P leaf P of P-allocation measured of use we compare efficient Thus, proportions to and history. aimed strategy life has contrasting life-history we with particular terms Therefore, a these exhibit environments. of context, to low-P meaning species broader allows the the that time, fractions to Over according respectively. and attenuata populations, 1981), crowded (Parry, in broadened ability competitive for Pate 2005, and Clarke, & Knox 2017, Pate, ek 0m lqo fbsllqi uretslto akn n otiig(e iormo ol:217.5 soil): of kilogram (per containing and dry P – lacking mass solution [(wet nutrient as liquid basal calculated for KNO of replicates was respectively, ten aliquot mg treatment gravel, were ml each limestone There 20 in sand. or A soils with week. laterite of filled to mass] of capacity were added dry layers layer Field was / other treatment. mm mass) substrate and each 100 kg surface, in soil 3.0 a species the treatment, treatment, each below soil SLIM mm each and 50 on For P SLAT added bags. based total was the plastic substrate imposed, For with The were lined pots. (SLIM). treatments were limestone the soil cylinder) + PVC Three sand tall and species. (SLAT), mm two laterite the + sand in sand was only, availability fractions than sand P (RGR) sand: foliar rate river growth washed among relative P greater of a allocation maintain and why types explore soil to designed was experiment 33 and 13 of Seeds method Sowing METHODS AND MATERIALS substrate. same N foliar higher ple oec o neeeyscn week. second every once pot each to applied CuSO mg 5 aki sessilis Banksia ° 9 ,115 S, 59’ K ° ’E.Tepoeac fthe of provenance The E). 3’ taeyt ecieslcinfrrpdpplto rwhi nrwe ouain n selection and populations uncrowded in growth population rapid for selection describe to strategy 3 aki attenuata Banksia ° 4m CaCl mg 74 ; vrawoeya,adtasiso frdainit h lshuews6%o aua ih.The light. natural of 60% was glasshouse the into radiation of transmission and year, whole a over C sa is 4 .5H ° 3 )uigarnoie opeebokdsg.Gasos eprtr utae between fluctuated temperature Glasshouse design. block complete randomised a using E) 53’ Total K 2 ;07m H mg 0.7 O; × taeit ed o nwtepyilgclpteno loaigPaogfla P foliar among P allocating of pattern physiological the know not do We strategist. hc xiisamr opportunistic more a exhibits which , P : 0% h oswr aee oacntn egto 0 ffil aaiytretmsa times three capacity field of 80% of weight constant a to watered were pots The 100%. > Total 2 SLAT 4 gK mg 140 ; ai n netmr nncecaisthan acids nucleic in P more invest and ratio .r and R.Br. 3 > BO LM(i.S,Shi S1, (Fig. SLIM 3 . gCoSO mg 0.5 ; 2 .attenuata B. SO tal. et 4 .sessilis B. 0m MgSO mg 80 ; .sessilis B. nd 90.MAtu isn(97 ondteterms the coined (1967) Wilson & McArthur 1990). , a,21.Acrigt h upir h ed of seeds the supplier, the to According 2016. May, edwsunknown. was seed 4 .7H .attenuata B. Kih)A .Ms .R hee(ucae from (purchased Thiele R. K. & Mast R. A. (Knight) sal ogo cosawdrrneo P-impoverished of range wider a across grow to able is tal. et 3 2 4 ;04m Na mg 0.4 O; .7H r 00.Tept 10m ne imtrx400 x diameter inner mm (100 pots The 2020). , lf taeythan strategy -life 2 ;2. gMnSO mg 28.9 O; and .sessilis B. .attenuata B. 2 .sessilis B. MoO .attenuata B. 4 .2H .attenuata B. san is 4 .attenuata B. 2 .H rw ihdffrn olP soil different with grown ;2 gFNET,was FeNaEDTA, mg 20 O; ycmaigteueand use the comparing by 2 ;1 gZnSO mg 10 O; r taeit while strategist, hngono the on grown when Banksia ol aea have would , and r .sessilis B. .sessilis B. strategy 4 .7H species 2 ° O; B. 18 Posted on Authorea 24 Jun 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159301657.74157566 — This a preprint and has not been peer reviewed. Data may be preliminary. ed htwr re 7@,4h)adwihdbfr acltn h vrg edwih.Rltv growth Relative weight. seed average the (ln calculating as before calculated weighed was and (RGR) 48hr) rate (70@C, ( dried weight were that Seed seeds LA. total (LA / M area DM mass leaf dry remaining (DM plant the leaves Total remaining calculate the to for dpi determined (DM 200 determined at LA was scanned (DM) + and mass (LA harvested dry -80 area were and leaf at plant calculate USA) York, stored to discolouration or dpi New and damage 200 “K”, visible at nitrogen Benchtop no scanned liquid with immediately leaves were in fully-expanded leaves 20 The submerged of plant. total each a from pots, harvested in were growing of weeks 50 After MD, Bethesda, NIH, Harvest 1.4, (ImageJ calculated 70 and at USA) CA, dried Beach, then Long were plant 1680, Leaves each (Epson USA). dpi of 200 leaf 1500 at mature of USA). measured One NE, density measurement. Lincoln, flux photosynthesis Inc., photon the LI-COR photosynthetic 400 (LI-6400, before a source day under light the measured LED on was red/blue watered a were using plants 2017 (P The 9th rate 7th, photosynthetic March net on harvest, 11:00 final the to Prior measurement Photosynthesis asprae,rltv rwhrt RR oncecai hshrs() oirntoe N n oirN foliar and leaf (N) concentration, nitrogen P foliar foliar (P), phosphorus total acid to nucleic fractions to P ANOVA (RGR) one-way rate foliar by growth of analysed relative were relationships area, types The substrate per intervals. across mass species confidence each 95% within means with in differences the while between means in differences The concentration. N Statistics area-based concentration to N rates area-based photosynthesis Leaf of concentration; ratio P the concentration area-based as concentration mass to calculated mass N rate Leaf P photosynthesis as leaf calculated of Total was ratio as the calculated as was culated concentration P GmbH, of Analysensysteme area-based combustion Elementar Leaf by Analyser, determined Combustion of was Macro sum (Vario digestion concentration sample acid the N Germany). by leaf is foliar confirmed dried Total Langenselbold, P was of assay. leaf P mg Pi Total leaf 30 by Total 1966). followed approx. fraction. (Ames, material, residual leaf method and ground blue metabolites described of molybdenum small in lipids, a as acids, using measured nucleic were (2004) Pi, procedures Golik above & the from Matusiewicz residues by by and extracts described in fraction method concentrations solubility residual Phosphorus differential a the and using metabolites) Pi. Yan leaves in other – powdered modified + P of as (Pi portion (2013), Yan Kitayama metabolites mg by & small 50 described Hidaka a lipids, (Pi) New in acids, P Metuchen, determined nucleic inorganic was SamplePrep, to determine to allocated Spex used P 2010, was The (Geno/Grinder sample powder mg fine 50 A a USA). to Jersey, ground were leaves Freeze-dried analyses nutrient Leaf weeks. in expressed respectively, harvesting, μ o mol mol 2 h eann evs tmadroswr eaae n re t70 at dried and separated were roots and stem leaves, remaining The . -1 h evsue o htsnhssmaueetwr ape,adtepoetdla area leaf projected the and sampled, were measurement photosynthesis for used leaves The . 2 DM = W 1 M .attenuata B. 1 DM + a esrduigfu oso 0( 10 of lots four using measured was ) 2 ° o 2ht esr r as(DM). mass dry measure to h 72 for C -ln W 2 2 n o tm lsros(DM roots plus stems for and ) DM + 1 × ( / ) M,adpooytei irgnueecec PU)was (PNUE) efficiency nitrogen-use photosynthetic and LMA, and tal. et ° T .Foe evswr reedidfrsvndy (VirTis days seven for dried freeze were leaves Frozen C. 3 efms e ra(M)wscluae sttlleaf total as calculated was (LMA) area per mass Leaf . .sessilis B. 2 21) eaoiePi endhr ssalmetabolite small as here defined is P Metabolite (2019). - n 4 T fatce evswsmaue ewe 00 and 10:00 between measured was leaves attached of ) 1 ,where ), ntesm usrt eeaaye by analysed were substrate same the on T μ o m mol 1 and 3 .attenuata B. -2 T .Ttlla M=DM = DM leaf Total ). 1 .Termiiglae neach on leaves remaining The ). s 2 -1 eetedtso oigand sowing of dates the were n CO a and ° o 2h h Mwas DM The h. 72 for C × M;PU a cal- was PPUE LMA; r3 ( 30 or ) 2 .TtlL LA = LA Total ). tal. et 2 ocnrto of concentration (2019). .sessilis B. 1 DM + t test, 1 2 ), 1 ) . Posted on Authorea 24 Jun 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159301657.74157566 — This a preprint and has not been peer reviewed. Data may be preliminary. LMwr oe hni lnsgoni adadSA Fg A,wietedffrne nmtblt P metabolite in differences the while 2A), of (Fig. of concentrations SLAT concentrations P and P lipid sand acid The in nucleic 2). grown plants (Fig. in in substrates concentration than all lower in were fractions SLIM residual and acid attenuata Banksia fractions phosphorus Leaf between PNUE in differences of for PPUE PNUE the while However, 2). ( (Table for substrate species PNUE any either in within PPUE species in sessilis two differences the substrate-dependent between no for rate were photosynthesis differences in substrate-dependent differences no no were there while SLAT, (P rates photosynthetic Net PNUE and PPUE rates, Photosynthetic N The SLIM. or SLAT in in ( that SLIM grown than or when higher SLAT species in N grown either The when (N for 1). than P (Table sand total types in foliar substrate grown to three N the total across foliar species of ratio concentration The 1D). (Fig. distinguishable not for was concentration N total attenuata foliar area-based for in not differences ( for substrate-dependent basis concentration also area P were an in foliar both on same for the expressed the substrates were basis, were across concentrations species P P both total when ( for foliar apparent SLIM mass-based basis in in mass grown differences a when The on than concentrations higher N and total SLAT, for leaf and basis The sand mass 1B). for a (Fig. concentration type on P substrate concentration total N foliar total Mass-based ( foliar 1A). substrates three (Fig. all type on substrate between differed concentrations any differences P for significant total concentrations no Foliar of were P (SLIM). leaves there limestone However, in + both 1A). sand relations for (Fig. and basis (SLAT) N mass laterite and a + P on sand sand, on in availability plants P growing of effect The concentrations nitrogen and phosphorus Leaf RESULTS USA). MA, Chicago, Northampton, Inc., Corporation, (SPSS by 19.0 (OriginLab analysed Statistics 9.5 were SPSS OriginPro coefficients the with correlation using graphed the performed analysis, and were regression US), analyses linear statistical by All determined T-test. were Student’s P acid nucleic to a infiatyhge hnta of that than higher significantly was .sessilis B. nalsbtae,teae-ae oirNcnetainwshge in higher was concentration N foliar area-based the substrates, all in .attenuata B. .sessilis B. .attenuata B. Fg D.I otatt ihrms-ae oirNcnetainin concentration N foliar mass-based higher to contrast In 1D). (Fig. and .attenuata B. .sessilis B. rw nsn a ihrta hto lnsgoni LM( SLIM in grown plants of that than higher was sand in grown .attenuata B. a h aei l usrt ye Tbe2.Mroe,teewr osignificant no were there Moreover, 2). (Table types substrate all in same the was n for ) .attenuata B. rw ntetresbtaetpswssand was types substrate three the in grown P .attenuata B. neeysubstrate. every in < .attenuata B. a ieetpten falctn efPt ii,mtblt,nucleic metabolite, lipid, to P leaf allocating of patterns different had .5:sand 0.05): .attenuata B. and .sessilis B. and .attenuata B. rw nSI eelwrta npat rw nsn and sand in grown plants in than lower were SLIM in grown > rw nSI eelwrta o lnsgoni ador sand in grown plants for than lower were SLIM in grown .sessilis B. .sessilis B. SLAT a ihrta for than higher was P 5 eehgetwe rw nsn,adlws nSLIM in lowest and sand, in grown when highest were < P > Total .5.Hwvr hr a osgicn difference significant no was there However, 0.05). Total naltresbtae ( substrates three all in o n usrt ye(al 2). (Table type substrate any for LM ncnrs ofla oa concentration, P total foliar to contrast In SLIM. < a rae hnta for that than greater was Total 0.05). P : P : .attenuata B. .sessilis B. Total P : .attenuata B. Total P Total < .attenuata B. ai o ohseiswslwrwhen lower was species both for ratio ai in ratio .5 i.1) oee,o narea an on However, 1C). Fig. 0.05, .sessilis B. a h aeptenfrtetwo the for pattern same the had ) > ( P SLAT and < and .sessilis B. P .5 al ) hr were There 2). Table 0.05, .attenuata B. .sessilis B. P > and .sessilis B. nalsbtae.There substrates. all in > < .5.Mroe,there Moreover, 0.05). .attenuata B. .sessilis B. LM(i.2) The 2B). (Fig. SLIM .attenuata B. .sessilis B. P .5 al ) The 2). Table 0.05, .attenuata B. < a significantly was .5 al 2), Table 0.05, a etdby tested was nfla total foliar in .attenuata B. nSA,or SLAT, in hnin than eealso were rw in grown neach in but , B. B. Posted on Authorea 24 Jun 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159301657.74157566 — This a preprint and has not been peer reviewed. Data may be preliminary. eeal erae ihicesn M Fg ) hsrltosi a o infiatfrlpdPand P P lipid in total LMA for and and significant P fractions not residual P between was the relationship relationship a of of This concentrations lack 5). The the m was (Fig. 2). in g Table LMA P acid 267 5, increasing nucleic (Fig. to with substrate 240 decreased growth from generally the varied Pi 5). on and area (Fig. residual depending concentration both acid, per concentration P nucleic P for total mass metabolite, of foliar concentration lipid, Leaf comparison with in P There pairwise positively P acid other 4B). concentration correlated any nucleic the all (Fig. in species, were and concentration detected both fractions concentration N were In concentration. N correlations foliar N other foliar and with No between 4A) 4C). correlation (Fig. (Fig. in concentration positive species of while P RGR a SLIM, The acid also in SLIM. nucleic was and lower SLAT foliar but both with SLAT, in correlated and equal and sand area lower in per but mass sand, same leaf the and was fractions, RGR P of foliar concentrations, RGR nutrient The foliar and RGR of relationships The for except attenuata substrates, all in 10% sessilis in that in than greater P acid lipid nucleic metabolites, in , P ( lipids total in of P proportion total The of proportions the in for difference Pi significant and no was There 3E). Fig. adta hngoni LTo LM( SLIM or SLAT for in Pi grown when than sand h rprin flpdPadncecai in P acid nucleic P and P foliar lipid total of to proportions fractions The P foliar of proportions The 2). otiue otelwrla N in leaf lower were lower fraction the to contributed P differed only N concentration The P 2E). residual (Fig. in SLIM The concentration not C). the but 2B, SLAT, with (Fig. SLAT, SLAT in in sessilis grown same B. when the species were of the but concentrations between SLIM, P and E). acid sand C, nucleic both B, and 2A, P (Fig. metabolite SLIM of The and concentration of SLAT concentrations P in Pi lipid grown and The plants P in acid those nucleic P, than metabolite greater P, were lipid The 2C). (Fig. SLAT ) hsrsl hw htmtblt n ifatoswr rvr o h bevddffrnei leaf in difference observed the for drivers were fractions (Table Pi substrates the and among P indistinguishable metabolite were fractions that other shows N the result for ratios This the 1). while SLIM, or SLAT P Total < .5 i.3) ovrey h rprino oa nncecai in P acid nucleic in P total of proportion the Conversely, 3A). Fig. 0.05, Total .attenuata B. P : a rae hnin than greater was Total a rae hnta of that than greater was Fg D.TeP ocnrtosof concentrations Pi The 2D). (Fig. .sessilis B. P : .attenuata B. Fraction ai for ratio .sessilis B. a rae o lnsgoni adta o lnsgoni LTo LM( SLIM or SLAT in grown plants for than sand in grown plants for greater was .attenuata B. aisfrmtblt n ifor Pi and P metabolite for ratios rw nayo h he usrts(i.3A). (Fig. substrates three the of any in grown .attenuata B. .attenuata B. h nyecpint eaiecreainbtenla rcinadLMA and fraction P leaf between correlation negative a to exception only The . a oe hnfor than lower was .attenuata B. .attenuata B. .sessilis B. Total nalsbtae ( substrates all in .sessilis B. hnin than ntetresbtae.For substrates. three the in P : .attenuata B. a rae hnta of that than greater was Total nSI Fg B ) h rprino eiulPwsbelow was P residual of proportion The C). 3B, (Fig. SLIM in .sessilis B. P rw nSA Fg D.Tepooto fttlPi iin Pi in P total of proportion The 3D). (Fig. SLAT in grown < naltresbtae ( substrates three all in .attenuata B. ai nsn.Terto fla N leaf of ratios The sand. in ratio .sessilis B. .5 i.3,C.Cnesl,tepooto fttlPin P total of proportion the Conversely, C). 3A, Fig. 0.05, -2 .attenuata B. P a rae hnta of that than greater was in 6 < o lnsgoni n ftetresbtae (Table substrates three the of any in grown plants for .attenuata B. .attenuata B. .attenuata B. .1;lkws,tepooto fmtblt in P metabolite of proportion the likewise, 0.01); nalsbtae Tbe1.In 1). (Table substrates all in eegetrta hs of those than greater were .attenuata B. eesgicnl oe o lnsgonin grown plants for lower significantly were .attenuata B. .sessilis B. .sessilis B. eesgicnl oe nsn hnin than sand in lower significantly were eelwrta hs of those than lower were .attenuata B. P n 1 o11gm g 131 to 119 and .sessilis, B. < .5 i.3E). Fig. 0.05, . l atosecp residual except factions P all , nSA n LM(i.2A). (Fig. SLIM and SLAT in .sessilis B. and .sessilis B. Total G a lohgetin highest also was RGR .sessilis B. .sessilis B. en oe hnta in that than lower being .sessilis B. P : nSA n SLIM and SLAT in .attenuata, B. Fraction a significantly was -2 rw nsand in grown a positively was in .sessilis B. nsn and sand in .sessilis B. nec P each in P < 0.05, the B. B. in Posted on Authorea 24 Jun 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159301657.74157566 — This a preprint and has not been peer reviewed. Data may be preliminary. h ieetfla -loainpten obndwt ieecsi M ewe h w pce reflects like (Clarke species Plants two fast requirements. grow the resource typically between and history LMA strategies life in history differences life with their combined in patterns differences P-allocation foliar different The fractions P and P- different traits Foliar on established become with and compared dpaw.wa.gov.au/). colonise limestone), http://florabase. to environment. over (FloraBase, the it or sand in allows laterite deep species presumably over two here (sand P the found soils of acquire concentration impoverished patterns to N distribution capacity leaf different higher higher the The This explain (Shi concentration. may P N SLIM higher acquire P-limiting the more to support capacity to higher P acid and nucleic species, in both P in more concentrations P acid attenuata that nucleic B. hypothesis and second in N our concentration leaf N supports with and correlated RGR strongly that was and RGR that (Kuppusamy found concentration We protein concentrations. in turnover. we N protein change since and However, hence rapidly synthesis. protein and not protein the of do rates which hence lower al. leaves, and and non-growing concentrations synthesis, mature rRNA protein lower studied abundance with of in ribosome consistent rate low concentrations is the A tested N the was LMA). leaf lower so decrease its lower hence to by and Therefore, evidenced expected species, as two be tissue, the sclerenchymatic can for in in investment similar in lower were fraction N a basis ‘extra’ acid of the area Therefore, nucleic an (PNUE). efficiency on the N-use expressed photosynthetic to concentrations P N leaf of and allocation greater with hlttela ocnrtosi ohseiswr o oprdwt h lblaeae(Reich average global the with compared low were in species higher both distinctly were in they P. concentrations 1991), thus, N and, leaf rRNA the for Whilst demand low a implies in which low, concentrations (Reich N environments other foliar from low plants of those in with presumably, compared low (Prodhan al. very concentration were et on N species results leaf differ both influence result of not Our concentrations did SLAT. the availability and substrate, P sand each in that grown showing in plants in species in two those concentrations than the N lower in Leaf of concentrations substrate. concentration P same total N carboxylate-extractable S1). foliar (Fig. greatest total gravel mass-based The foliar limestone similar SLIM. mass-based in the in was to lower lowest contrast 35% the In about and and sand, sand, in concen- was in P concentration total greatest foliar P were mass-based species of patterns both same of the found both We substrates. for three tration the of any in concentration upre.A motn nigwsta h rabsdfla ocnrtoswshge in higher was concentrations P foliar area-based the concentrations that P was ta total finding foliar important mass-based An the availability, supported. P soil decreasing both with of that was hypothesis first concentrations Our nitrogen and phosphorus total Foliar DISCUSSION hnin than 04,tefse aeo rti ytei uthv enblne yfse aeo rti breakdown, protein of rate faster by balanced been have must synthesis protein of rate faster the 2014), , 91,wihrflcstelwfla RAcnetainin concentration rRNA foliar low the reflects which 1991), , .attenuata B. .sessilis B. (Shi .sessilis B. .attenuata B. tal. et and naltresbtae,btteewsn infiatdffrnei asbsdfla P foliar mass-based in difference significant no was there but substrates, three all in 00,wl aeahge oirN foliar higher a have will 2020), , ae ntesmlrsz fterncecai ol.I u td,terelatively the study, our In pools. P acid nucleic their of size similar the on based , .sessilis B. and .attenuata B. .sessilis B. .sessilis B. .sessilis B. tal. et ol eraei epnet erae viaiiy n hswas this and availability, P decreased to response in decrease would .sessilis B. tal. et .attenuata B. .attenuata B. 03 n rdc ed eoetenx catastrophe, next the before seeds produce and 2013) , .sessilis B. and rw naltresbtae;tefla oa concentrations P total foliar the substrates; three all in grown 00 in 2020) , hnin than hc xiisamr potnsi rwhsrtg than strategy growth opportunistic more a exhibits which , eesgicnl rae hntoein those than greater significantly were .sessilis B. 7 .attenuata B. a lottieta of that twice almost was .sessilis B. oprdwith compared and Total niaeta rti ocnrtoswr very were concentrations protein that indicate .sessilis B. .sessilis B. P : hnin than .attenuata B. .sessilis B. Total h ihrNcnetaincorrelated concentration N higher The . oee,rtso photosynthesis of rates However, . rw nSI eeapo.20% approx. were SLIM on grown .sessilis B. ai than ratio .attenuata, B. .attenuata B. tal. et nams ai a reflection a was basis mass a on ae prostrata Hakea .sessilis B. (Sulpice .attenuata B. 06.Tefla oa N total foliar The 2016). , .attenuata B. ntetresubstrates three the on hc srsrce to restricted is which .attenuata B. hngoni the in grown when ihan with tal. et rw nthe in grown 04,and, 2014), , (Proteaceae) .attenua- B. r n invest and i.e. selection tal. et r or fire This . et , Posted on Authorea 24 Jun 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159301657.74157566 — This a preprint and has not been peer reviewed. Data may be preliminary. napcso hswr,adRbCes n ilPaiifrhl ihmitiigtepat nteglasshouse. supported the was in research plants This the (201508220115). maintaining grant with help Project for Council assistance Scholarship Piasini their China Bill by for and Hayes supported Creasy E. was Rob Patrick Z.H. and and work, this Prodhan of Asad Wang, aspects Yunhe in Ding, growth Wenli Ilyasova, in Albina thank differences We their match and availability two the P ACKNOWLEDGEMENTS of soil traits to that foliar response surmise distinct concentrations We in P and strategies. strategies species. foliar patterns reduce two adaptive nutrient-allocation can different the foliar plants reveal why of The explaining soils. distributions basis P-poor different functional on the the likely for are account patterns than protein P-allocation may ratios greater and N:P for and needed soils, acids likely P-impoverished nucleic is to which P synthesis, of protein greater that observations support The may turnover. acid nucleic limitation. in investment P under leaves the Both 6). (Figure found are they and CONCLUSIONS availability which P in soil niches to ecological response the in define allocation may P in that of (Veneklaas LMA patterns strategy life-span different lower growth revealed leaf with a species shorter associated two and and the LMA P RGR of a high lipid traits higher with with foliar a of plants line with proportion slow-growing in (Villar associated lower in to partially LMA traits a greater was than low P, is finding Pi) with P acid This plants + structural nucleic alone. of fast-growing P sand concentration in with (metabolite the compared than P that availability showed P metabolic that lower to study the P to addition, of response In in proportion here. greater however, showed tested; have a substrates we of as allocation P, the structural by sustained is of for that metabolites in than P for total those of slow-growing than proportions the higher in and as concentrations P Pi, 1995). (G metabolite accumulate (Mimura, vacuoles, The ones than in decreases rather fast-growing Pi availability P, in of P organic accumulation than growth-sustaining as the into species. species upon reflect Pi slow-growing reservoir generally convert drawn in a supply species be as growing Pi greater serve then with typically vacuoles concentration can is Cell P which SLIM. which foliar in plants, total grown most in when in than Changes higher Pi slightly excess SLAT, and for sand in slow-growing grown with in when compared concentration roots Pi deep Pate The to 2005, biomass Clarke, more & allocate Knox to 2017, ability Pate, the as well as sessilis P, Pate acid 2005, nucleic Clarke, in investment & Knox Pate 2017, Pate, 2011, & Lamont, Unlike & quickly. (Groom cycle provide life (Shi lignotubers may the seeds capacity complete larger turnover, ( synthesis and environments and with fluctuate) protein changing synthesis may high availability and protein water A variable rapid where 2012). to support (Raven, acclimate to proteins to ribosomes, flexibility damaged thus, of and, replacement rRNA Pate including P-rich 2005, Clarke, in & investment Knox greater 2017, Pate, & (Bowen drought .attenuata B. (Shi tal. et .attenuata B. 00.Tu,i osntne ogo atadcmlt t iecceqiky(oe & (Bowen quickly cycle life its complete and fast grow to need not does it Thus, 2020). , and .attenuata B. tal. et .sessilis B. rw nalsbtae.Hdk iaaa(09 ugse hthg PPUE high that suggested (2009) Kitayama & Hidaka substrates. all on grown .attenuata B. 00 n ihrLA a h blt orsru rmeiomcbd or buds epicormic from resprout to ability the has LMA, higher and 2020) , aki sessilis Banksia .attenuata B. xiie nqepteno loaigPt ieetPfatoswithin fractions P different to P allocating of pattern unique a exhibited o lnsgoni LM oevr h PEof PPUE the Moreover, SLIM. in grown plants for tal. et .attenuata B. a ihrlpdPcnetain hngoni LTadSLIM and SLAT in grown when concentrations P lipid higher had tal. et .sessilis B. tal. et a ihrLAadlwrNcnetain PE allocation PPUE, concentration, N lower and LMA higher a had 1990). , 90.Tu,slcinin selection Thus, 1990). , loae oePt uli cd than acids nucleic to P more allocated 91,asrtg soitdwt lwrRR(Bowen RGR slower a with associated strategy a 1991), , tal. et .sessilis B. r osbyaatv rist rwhi oeseverely more in growth to traits adaptive possibly are a ihrta hti h faster-growing the in that than higher was 8 tal. et 06.I te od,agetrpooto of proportion greater a words, other In 2006). , a oe M than LMA lower a had 90.Ti taeymyrqierelatively require may strategy This 1990). , i.e. .sessilis B. hlo adoe aeieo limestone, or laterite over sand shallow .attenuata B. tal. et hnin than 02.Oeal h unique the Overall, 2012). , .sessilis B. swl,20) hs fast- Thus, 2004). usewell, ¨ .sessilis B. .sessilis B. a ae nalower a on based was .attenuata B. .attenuata B. .attenuata B. eesignificantly were Banksia , .attenuata B. a greater was .sessilis B. species r all are This . nall on B. Posted on Authorea 24 Jun 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159301657.74157566 — This a preprint and has not been peer reviewed. Data may be preliminary. ae .. ulem eer . ld ..&LmesH 21)Climehne hshrstxct in chronosequence. toxicity Bay phosphorus Jurien Calcium-enhanced the (2019) along H. Proteaceae Lambers soil-indifferent & and P.L. calcifuge Clode C., Pereira Guilherme P.E., Hayes 164 G ro ..&Lmn ..(01 einladlcleet nrpoutv loaini pcri and epicormic in allocation reproductive on effects local and Regional bioche- of (2011) its populations and B.B. lignotuberous microalgae Lamont marine & in P.K. C:N:P P- Groom of variability of revisited: Effect Redfield (2003) (2002) ( basis. J. P.K. tomato Roche mical Mohapatra of La diameter & & stem R. J.J. Geider and Adu-Gyamfi fruit K., in changes Ohkura rhythmic J., esculentum and copersicon Ito partitioning in K., photoassimilate photosynthesis on Lei on deficiency nutrition M., phosphorus Okada of K., Effects Fujita (1990) N. Terry & Sunder- I.M. max Sinauer, Rao Glycine Perspectives. T.K., and Raab Principles A.L., Plants: Fredeen of Nutrition N:P Mineral and (2005) MA. history, A.J. land, Bloom life & size, E. Organism Epstein processes. (1996) ecosystem J.H. and cellular Schampel of & 684. view now. N.A. unified is MacKay a future Toward D.R., stoichiometrystoichiometry: The Dobberfuhl biology: J.J., new Elser the and Stoichiometry (2007) phosphorus. A. 1404-1405. of Hamilton re-mobilization & and J.J. use Elser the Environment in and efficiency Cell extreme Plant, (2007) exhibit H. soils Lambers phosphorus-impoverished & CO F.M. ely Freimoser for E.J., requirements Veneklaas phosphorus M.D., in after Denton Increases persistence (1990) drive E.W. K.J.E. Barlow species. resources Knox & pine and M.L. & enriched Reed protection N.J. P.J., Milham buds, Enright J.P., How G., Conroy Burrows trait: F., functional Ojeda key B.B., a fire. Lamont as J.J., Resprouting Midgley (2013) M.J., autumn Lawes and P.J., fractions trees. phosphorus Clarke taiga and nitrogen deciduous in and changes evergreen Seasonal (1983) in R.A. retranslocation Kedrowski & F.S. photosynthesis III of Chapin response the on leaves. nutrition spinach phosphorus in of of 3-phosphoglycerate Effects taproot (1988) S.C. young CO Wong the to & K.C. in Woo distribution A., evolution. starch Brooks homoplastic and of evidence tissue Possible storage (Juss.): Proteaceae of Ecology Australian Austral Patterns of resprouters (2017) Complex and J.S. seeders VIII. obligate Pate vol. & phosphatases. B.J. Bowen and eds. phosphate V, total Ginsburg phosphate, FN, USA. inorganic Elizabeth of In: Assay carbohydrates. (1966) B.N. 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No reuse without permission. — https://doi.org/10.22541/au.159301657.74157566 — This a preprint and has not been peer reviewed. Data may be preliminary. ec .. h . atr ..&ElwrhDS 19)La iepna eemnn fla structure leaf of determinant a as lifespan Leaf species. (1991) tree D.S. amazonian and Ellsworth 23 growth & among ratio, M.B. function DNA Walters and : C., RNA Uhl among P.B., Relationships Reich (2010) nitrogen C.E. trees. Lovelock to mangrove & in relation I.C. stoichiometry in Feller elemental M.C., requirements Ball phosphorus R., and Reef RNA plant partitioning and enzymes. cycle turnover carbon Calvin and fixation. Protein and photosynthesis, (2012) exchange, status, gas J.A. phosphate growth, Raven in Leaf Changes (1989) I. N. beet: Terry sugar & in A.L. Fredeen ni- of I.M., control Rao environment. Tight phosphorus-impoverished (2016) extremely P.M. an Finnegan Environment in & evolved and that H. species Cell Lambers plant R., a Hoefgen in M., acquisition Watanabe trate R., oligo- Jost related M.A., and Prodhan laterite inter-relationships? Proteaceae, causative of or Co-occurrence 529-560. associations (2001) P.D. coincidental Galloway soils: & trophic W.H. Verboom characteristics J.S., Australia. storage Pate S.W. and of growth ecosystems Seedling (1990) Mediterranean-type J. of Kuo species 585-601. & resprouter A. and Hansen seeder species-rich B.J., of the Bowen to R.H., Froend concept J.S., mimic Pate ecosystem the of Australia. Application Western (1999) of T.L. lands Bell & J.S. fire-sensitive Pate of ( characteristics Restionaceae morphological of and species growth Western-Australia. (resprouter) Contrasting western fire-resistant (1991) and K. Dixon seeder) & (obligate K. Meney J., Pate hn ..&LmesH 20)Cutrros uist ncontext. in curiosity A roots: Cluster (2005) H. Lambers & ( M.W. rice Shane of quality and yield grain Growth, CO (1997) elevated J.P. Conroy & S.P. Seneweera icraRse ..&BiEN 21)Anwdtie a fttlpopou tcsi utainsoil. Australian in stocks phosphorus total of map detailed chemical new Environment A and among Total (2016) costs differences the E.N. construction to of Bui Science compared in & R.A. as Differences Rossel small (2006) Viscarra are H. species Poorter woody & evergreen Y. and families. Jong deciduous De between J.R., in composition Scheible efficiency Robleto phosphorus-use C.A., R., improving Price for Villar W.C., Opportunities Plaxton (2012) C.E., J.A. Raven Lovelock plants. & P.M., P.J. crop Finnegan White J., M.W., density Shane Bragg wood W., H., increase Lambers limitation phosphorus E.J., Veneklaas does & Why M. (2006) seedlings? J.P. Stitt grandis Conroy Eucalyptus E., & in Veneklaas K.D. F., phosphorus- Montagu photosynthetic Jost Teste D.S., high S., W., very Thomas the Arrivault Scheible for species. 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Princeton, Press, University Princeton . 505-525. , 16-24. , 24 37 1064-1072. , 1276-1298. , and ln n Soil and Plant utainJunlo Botany of Journal Australian hmshr restiads Shemisphere . .sessilis B. rz sativa Oryza ln Physiology Plant ihcnrsiglife contrasting with , naso Botany of Annals 274 . nrsos to response in L.) 101-125. , 43 Banksia rmsouth- from ) 90 1131-1136. , 814-819. , Plant, wood- 49 65 , , Posted on Authorea 24 Jun 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159301657.74157566 — This a preprint and has not been peer reviewed. Data may be preliminary. usrts( substrates relative ** means the for are (RGR) affects rate Values growth efficiency factor relative P-use and the photosynthetic fraction, that containing PPUE: area, indicates per arrow 1 mass dashed Table leaf red LMA: phosphorus, factor; P: another rate. affect growth that factors seeder indicate the and (a) i.6 Fig. limestone BA: plus plant. sand individual an represents for (right) symbol 1. (LMA) Table area and per 2, mass and 1 5 Figures Fig. in found be can data Individual eaoi P Metabolic / N Leaf P 100 Lipid / N Leaf Pi / N Leaf P leaf Total / N Leaf Items for P acid nucleic to N foliar and (N), 4 Fig. limestone * significant, not means ns means are Values (C), 3 Fig. ( substrates among species two * significant, means are concentrations (Pi) P 2 Fig. limestone. plus by sand SLIM: species laterite, two the between P differences significant show asterisks phosphorus of foliar trations of Responses 1 (2019) chronosequence. Fig. P.M. dune Finnegan Myr & 2 H. a legends along Lambers Figure diverse J., are Pang age Z., soil Han to fractions X., Zhang L., Yan ieecsfrtetoseisaogsbtaetps( types substrate among species two the for differences < P eiulP residual .0.Dffrn etr ersn infiatdffrne mn usrts( substrates among differences significant represent letters Different 0.001. < igassmaiigfla risaetn eaiegot ae o h resprouter the for rates growth relative affecting traits foliar summarising Diagrams eainhp ewe oirpopou P rcin n oa oirPcnetain(et,adleaf and (left), concentration P foliar total and fractions (P) phosphorus foliar between Relationships h ii hshrs(P) phosphorus lipid The efms-ae oa P total mass-based Leaf h rprin fttlfla hshrs()a ii P lipid as (P) phosphorus foliar total of proportions The eainhp frltv rwhrt RR ofla uli cdpopou P,fla nitrogen foliar (P), phosphorus acid nucleic foliar to (RGR) rate growth relative of Relationships 0.01, asrto o oa oirntoe N ottlfla hshrs()adt nec oirP- foliar each in P to and (P) phosphorus foliar total to (N) nitrogen foliar total for ratios Mass . ± aki attenuata Banksia P *** P E( SE < < (D) adSA LMSn adSA SLIM SLAT Sand Sand SLIM 119 159 SLAT 27.1 Sand attenuata Banksia P .5.SA:sn lsltrt,SI:sn lslimestone. plus sand SLIM: laterite, plus sand SLAT: 0.05). .5 ** 0.05, n ± ± < ± ± ± ± n nrai (Pi) P inorganic and aki sessilis Banksia 0;atrssso infiatdffrne ewe h w pce by species two the between differences significant show asterisks 10); = 0b168 168 b 10 161 a 25 b 14 E( SE E( SE . 33.4 b 1.8 .0.Dffrn etr ersn infiatdffrne o h w pce mn the among species two the for differences significant represent letters Different 0.001. (E) n n P nlae of leaves in 0;atrssso infiatdffrne ewe h w pce by species two the between differences significant show asterisks 10); = 0;atrssso infiatdffrne ewe h w pce by species two the between differences significant show asterisks 10); = P < and P .1 *** 0.01, attenuata Banksia < (A), aki attenuata Banksia (A) < .5 ** 0.05, ± ± ± .sessilis B. ± .5.SA:sn lsltrt,SI:sn lslimestone plus sand SLIM: laterite, plus sand SLAT: 0.05). 6a177 158 a 36 176 a 27 a 46 . 33.1 a 2.5 b.Sldlnsidct oncin ewe atr;dse arrows dashed factors; between connections indicate lines Solid (b). eaoieP( P metabolite , n concentrations N and aki attenuata Banksia aki attenuata Banksia P (E) P < .attenuata B. < for .0.Dffrn etr ersn infiatdffrne o the for differences significant represent letters Different 0.001. rw ntresbtae.Vle r means are Values substrates. three in grown attenuata Banksia .1 *** 0.01, aki attenuata Banksia ± ± ± ± and 8a189 294 a 38 253 a 23 a 46 12 . 45.9 a 3.6 B P aki attenuata Banksia .sessilis B. ,ncecai P acid nucleic ), < BS: , and P and .5.SA:sn lsltrt,SI:sn plus sand SLIM: laterite, plus sand SLAT: 0.05). (B), < .sessilis B. b attenuata Banksia .sessilis B. .sessilis B. .0.Dffrn etr ersn significant represent letters Different 0.001. *** t n rabsdP area-based and ± ± ± ± (A) rw ntresbtae ( substrates three in grown and test. 6b 76 b 86 9b 29 8.9 e Phytologist New .sessilis B. eaoieP( P metabolite , *** *** *** ns ausaemeans are Values . (C), and LT adpu aeie SLIM: laterite, plus sand SLAT: , rw ntresbtae.Values substrates. three in grown o significant, not .sessilis B. b 45.9 sessilis Banksia 294 253 189 eiulP residual P *** ± ± ± rw ntresubstrates. three in grown < ± 6b 76 b 86 9b 29 8.9 (C) .5.SA:sn plus sand SLAT: 0.05). 223 *** *** *** B ntresubstrates. three in n N and (D) aki attenuata Banksia ** 1621-1633. , ,ncecai P acid nucleic ), ± ± P n a 63.2 a 436 390 sessilis Banksia 274 n inorganic and t E( SE E( SE *** *** (D) 0.Each 30). = < et snot ns test. ± ± ± ± .1 *** 0.01, n n 127 a 67 1a 41 7.4 concen- 10); = 10). = t t test. test. *** *** 56.9 a ab 398 356 sessilis Banksia 239 *** *** ± ± ± ± 95 a 80 4a 54 7.72 *** ** Posted on Authorea 24 Jun 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159301657.74157566 — This a preprint and has not been peer reviewed. Data may be preliminary. and-adaptation-to-low-phosphorus-habitats allocation-patterns-in-banksia-attenuata-and-banksia-sessilis-differing-in-growth-rate- Figures.docx file Hosted m ( rates Photosynthesis to (P) phosphorus acid nucleic by of species ratio two foliar the between the 0.01, differences and significant (LMA) show area asterisks per of mass P foliar organic (PNUE), efficiency use 2. Table g (mg RGR P Lipid / P acid Nucleic P Residual / N Leaf P acid Nucleic / N Leaf Items g ( PPUE ( s ( PNUE oogncP organic to P acid nucleic of ratio The m (g LMA P μ -1 -1 -1 -2 o g mol ) < week s P s *** -1 .5.SA:sn lsltrt,SI:sn lslimestone. plus sand SLIM: laterite, plus sand SLAT: 0.05). μ -1 ) -1 mol μ P -1 ) mol htsnhssrts htsnhtcpopou-s ffiiny(PE,pooytei nitrogen- photosynthetic (PPUE), efficiency phosphorus-use photosynthetic rates, Photosynthesis ) N < -2 240 ) aki attenuata Banksia .0.Dffrn etr ersn infiatdffrne o h w pce mn h substrates the among species two the for differences significant represent letters Different 0.001. vial at available 95 1.52 416 105 attenuata Banksia adSA LMSn LTSLIM SLAT Sand SLIM SLAT Sand attenuata Banksia 15.3 234 8.9 0.355 ± ± ± . 93 a 2.6 ± ± 4a447 a 94 114 a 9 ± ± .8a1.48 a 0.28 ± . 7.9 a 2.2 3a264 a 53 5b248 b 15 ± 15.4 a 3 .2 0.378 a 0.026 https://authorea.com/users/336556/articles/462215-foliar-nutrient- attenuata Banksia and ± ± ± . 86 a 3.7 ± attenuata Banksia .sessilis B. 4a375 a 94 108 a 11 .5a1.46 a 0.25 ± ± ± ± . b6.7 ab 1.0 8a223 a 38 1b267 b 21 ± . 10.8 a 1.3 .4 0.382 a 0.042 attenuata Banksia rw ntresbtae.Vle r means are Values substrates. three in grown ± ± ± . 123 b 4.6 ± 6a692 a 66 136 a 13 13 .6a2.19 a 0.16 attenuata Banksia ± ± ± ± . 8.6 b 1.0 2a377 a 52 4a119 a 24 ± . 13.4 b 1.8 .2 0.420 a 0.023 a a attenuata Banksia ** *** t ± ± ± test. ± 195 b 29 a 4 0.44 ns *** sessilis Banksia a ** ** o significant, not ± ± ± ± . a 2.4 64a 86.4 b 7 ± . a 2.2 0.044 sessilis Banksia 2.19 a 692 136 123 a *** ** ** ns ± ± ± ns ± *** 195 b 29 a 4 0.44 *** ** * 13.2 sessilis Banksia 506 7.9 121 a 0.411 ns P ± ± ± ± < ± . a 3.5 607 186 sessilis Banksia b 115 a 2.35 6 a 269 b 9 ± E( SE . a 6.2 *** 0.05, *** 0.043 ± ± ± ** ± ns n 8a 98 a 28 7.6 * ns 0.55 ** 10); = P ** *** < sessilis Banksia 10.7 410 7.3 131 a 0.434 ns a 772 ab 159 sessilis Banksia b 110 a 2.53 ± ± ± *** *** *** ± . a 1.9 *** 1 a 116 a 6 ± . a 2.3 ± ± ± ± 0.041 258 25 6.1 ** 0.58 ns *** ns