Posted on Authorea 25 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158515652.24058515 | This a preprint and has not been peer reviewed. Data may be preliminary. islcfiaini ln cusi ue n elwlsbtas netaellraditrellrspaces Exley intercellular & (Law and deposition silica extracellular for in “catalyst” a 1985; Franklin also be & could but Dayanandan Callose Kaufman, 2019). walls 1985; Hodson Parry 2001; cell Tubb & & thus Sangster and Hodson Hodson, Sangster, and lumen 1959; agent)”, Kitagishi in & cellular Onishi occurs active (Yoshida, functions. plant Si-induced an Si on to defines in deposits opposed model Biosilicification silica (as Their extracellular plant stresses hypothesis”. of and obstruction against role 2016) agent the “apoplastic Zhu highlights prophylactic the & “extracellular through Zhao an plants Coskun (Kang, as in 2016). efficiency functions Datnoff photosynthetic Si-induced & increase the they Babu (Tubana, while Verheggen Corn´elis & 2013) biomass Walgraffe, Tombeur, Junior de Leroy, Gomes Davis B´elanger 2006; Cai & R´emus-Borel, & 2005; & Hartley (Fauteux, Menzies McNaughton diseases & Tarrants, and Massey pest (McNaughton, 2006; 2019), herbivores Meyer against & protection Keeping progress alleviate plant 1985; indeed further enhance plant stimulates 2017), in recognition functions Dudel Si-induced This & (Adrees practices. stresses substance. have agricultural advances beneficial in abiotic of Si number of various of a status exploitation 1994), (Epstein the optimal biology the to plant towards in Si (Si) elevate silicon to of anomaly contributed the of recognition the Since Introduction leaf Soil as cellulose correlated. and through negatively silica Si between were leaf counterbalance concentrations increasing the Si with and and increased biosilicification on cellulose deposits of depending Ash-free silica magnitude silicified, components. of the fully structural cells. size veins markedly deposits prominent The silicified impacted silica in climate conjoined Epidermal highest. arranged and of cells or transpiration. short number lowest plant and was Soest increasing and long Van concentration soil an to the Si in by expanded leaf tropical Si determined or digestion. bioavailable the three were cells wet leaves to whether in silica through in related to collected leaves biopolymers was limited leaves from C-rich struc- leaf either sugarcane extracted The were as in ESEM-EDX. physically in accumulation biopolymers by were situ Silicon accumulation Si C-rich deposits in for silica and silica method. deposition mapped affect silica Plant and silica climate between observed climate. studied and were balance and we Leaves Soil and soil Here, patterns in environments. biosilicification differing known. and environments on poorly species roles plant is Their of components variety tural species. a plant in given effects in beneficial has (Si) Silicon Abstract 2020 5, May 4 3 2 1 Comp`erePhilippe ´ lxd Tombeur F´elix de and patterns balance. silicification silica-cellulose foliar affect climate and Soil nvri´ eLi`ege agronomiques Universit´e de Recherches de wallon Centre Louvain de Universit´e catholique Liege of University 1 4 hre adrLinden Vander Charles , n rn Delvaux Bruno and , tal. et 05 ok esmn21;Meunier 2016; Leishman & Cooke 2015; 2 2 enToa Corn´elis Jean-Thomas , tal. et 1 21)pooe opeesv oe linking model comprehensive a proposed (2019) tal. et 1 rn Godin Bruno , tal. et 08 aag,Amorim Camargo, 2008; 07 e,Schaller Neu, 2017; 3 , Posted on Authorea 25 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158515652.24058515 | This a preprint and has not been peer reviewed. Data may be preliminary. h ocnrtoso iado ells (Schoelynck cellulose between of relationship Epstein inverse and 1983; The al. (Raven Si structures. component Elbaum vegetal of of structural Bauer, concentrations resistance compression-resistant 1999; mechanical the a the (Epstein as to tissues act contributing hence plant indeed 1994), may strengthens (Ando, Silica which plants 2011). in 2000), Weiss Keeling-Tucker silica & of & role Li Perry structural 2002; 1991; The Ajiki Fraser 2018). Exley & & Suzuki Moore Fujii, Stokes, Kakuda, Law, Guerriero, 2015; Exley 2011; akn feiec o h tutrlrl fS n hsblnewt -ihbooyes(Bauer is Soukup biopolymers literature C-rich 2016; However, (H with Struyf acid balance 2017). & thus Monosilicic Hartley Schoelynck and 2016; & Si Hartley of Osborne & role Schaller Rees, DeGabriel structural 2012; Wade, Cooke, the Dudel for Simpson, & evidence 2016; of Brackhage Johnson lacking Schaller, & 2012; Allsopp Leishman Sallam, & (Cooke plants in oArli l ie.Tewtetst steAdsloe(al )weetediyEPmaue during measured ETP daily the where 1) (Table one Andosol the December is from is site decreases MAT wettest precipitation mm, The Monthly 1275 1a). sites. to (Fig. all amounts July in MAP to April December site, Vertisol from to mm the 100 In 25.4–25.3 below limestone 1). and invariably (Table mm Pleistocene mm 2910–3170 from respectively 34.2–34.5 26.7 are, derived is 1a) ETP materials Fig. ten-day 1, smectitic relative (Table MAT in and formed MAP sites, Vertisol and weathered The highly 1965). is (Komorowski Lagache Nitisol The & Komorowski 2014). 1965; Daage Boudon (IUSS Lagache BP; system & ka WRB (Colmet-Daage ash the (30–18 andesitic in old Vertisol from and formed Andosol Nitisol, as out (16 ( Guadeloupe sugarcane in in out carried was fieldwork Our setting Environmental role compensatory leaves methods mechanical that a and hypothesize as Materials concentrations further We cellulose the accumulation. higher of have Si that magnitude will of the hypothesize concentration (Yamamoto level of We Si contrasted drivers foliar key via conditions. low be epidermis climate with will leaf potential and on evapotranspiration structural deposits soil and leaf silica minerals contrasting as weatherable biopolymers in soil C-rich of and cultivated reserve Si between sugarcane balance in with and patterns balance components the biosilicification the of the for study thus analysis we and literature. an Here, silica in Finally, of unclear accumulation. conditions, role remains Si mechanical natural which C- the plant under biopolymers, for and controlling investigated C-rich support Si factors be provide between to the could tradeoff deserves patterns of biosilicification it a understanding literature, how since complete understanding in addition, a regulation, highlighted In with stress been important. and has is functions biopolymers process plant rich on this weatherable silicification impact of foliar conditions reserve of environmental effect the beneficial is the property Given soil 1967; H Handreck important of source & An primary (Jones 2005). (Henriet the conditions represents Schwab climatic which and & minerals, 2014), Banks primary al., Anderson, et Benke, & (Li Dorsey, Quigley properties Tombeur, 1994; Euliss, water de Weiner, 2016; and & physico-chemical Anderson Broadley & Rosen soil Donati Jaeger, including 2019; & Quigley, De 2020), 2016; Henriet, Mead Cornelis Delvaux 2008a; & & Lalibert´e, White, Delvaux Cornelis Lambers Turner, & 2008b; (Hodson, Draye Delvaux Bodarw´e, Nahon Dorel, & variation Henriet, & Opfergelt 1999; Rouiller Luiz˜ao, phylogenetic Dorel, Alarcon Chauvel, (Lucas, & on properties Colin and depends Meunier, processes 1993; plants (SiO soil B´elanger 2016), vascular opal-C & Deshmukh amorphous in 2005; as uptake precipitation Si silica promotes The loss water where 2006) 07 Suzuki 2017; , ° hlttema eaietndyEPrahs4. m n h vrg otl rcptto is precipitation monthly average the and mm, 42.0 reaches ETP ten-day relative mean the whilst C tal. et tal. et tal. et 08 ;Klotzb¨ucher b; 2008a tal. et ° 02 urir,Humn&Lgy2016). Legay & Hausman Guerriero, 2012; 5N 61 15’N; 4 05 ne re odtos(omtDae&Lgce16) nteNitisol-Andosol the In 1965). Lagache & (Colmet-Daage conditions drier under 2005) SiO 02 Yamamoto 2012; , tal. et 4 stknu rmsi ouin rnlctdt ln rnprto ie (Ma sites transpiration plant to translocated solution, soil from up taken is ) ° 3W.Sislreydffrd(al ;Cle-ag aah,16) hykey they 1965): Lagache, & Colmet-Daage 1; (Table differed largely Soils 33’W). 97 eeoe nEcn neii s nprui odtos(Colmet- conditions perhumid in ash andesitic Eocene on developed 1987) , tal. et tal. et tal. et 2015). acau officinarum Saccharum 05 satiue otehrns rpryo iia(er & (Perry silica of property hardness the to attributed is 2015) 02 ihihsteblnebtenS n components C and Si between balance the highlights 2012) , 2 tal. et 4 00 rlgi Bnla 01 Klotzb¨ucher 2001; (Bonilla, lignin or 2010) , SiO 4 nsis n hso iaalbefrplants for available Si of thus and soils, in . xliain salse ntresites three in established exploitations L.) tal. et tal. et 07 az2019). Katz 2017; 2 .nH 05.TeyugAndosol young The 2005). tal. et 2 )frigphytoliths. forming O) ° hra h mean the whereas C 09 rw Powell, Frew, 2019; tal. et 2011; tal. et et Posted on Authorea 25 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158515652.24058515 | This a preprint and has not been peer reviewed. Data may be preliminary. TB nsiswscmue stesmo ao laieadakln-at ain C,M,KadN in Na and K Mg, (Ca, cations alkaline-earth and alkaline major of sum the the of as dissolution computed cmol After was 1992). soils Sanzolone in HNO & (TRB) 10% (Chao in in concentrations Li-metaborate bead elemental and fusion Total Li-tetraborate ICP-AES. 450 with by Sommer at crucible concentration calcination & graphite after Si CaCl Herrmann a determined for Conley, of were analyzed Saccone, ml soil and Sauer, 40 filtered with 1975; shaken was Chapman were supernatant 1 soil & H by of (Haymsom in determined grams Four CaCl soils were measured 2006). 1982). in (CEC) was Page capacity Si & pH bioavailable exchange Sommers Soil cation (Olsen, assess mm. pH7 and acetate cations 2 ammonium Exchangeable at M 1g:5ml. sieved of and ratio air-dried lid:liquid a were constitute samples to Soil collected randomly were was analysis subsamples midrib sugarcane Soil leaf The dewlap) twenty avoided. visible and were (top suckers subsamples sample. TVD and soil composite The tillers three 2017. while replicate, April stalks each in or depth shoots (McCray soil primary removed cm from 0–20 sampled at were leaves sampled were topsoils The sites between collection differed Sample cultivars kg sugarcane g The (<0.02 cultivated. cultivars the between been in long ETP 1). increasing has (Table and sugarcane availability environments, water these decreasing In soil to in leading minerals clay differences 1), Andosol swelling (Table from Si-rich Andosol–Nitisol–Vertisol. the originates regime secondary sequence contrast moisture in which mineralogical soil gibbsite in This & Vertisol and and 1965). (Churchman the age Lagache substances desilication in & retained strong allophanic was (Colmet-Daage denoting Al-rich silica accumulate 2003), contrast, Nitisol, Delvaux In & the 2012). Ndayiragije in Lowe 1965; were oxides Lagache bases of Fe and & processes and silica (Colmet-Daage the threshold, kaolinite enhanced agreement that as leaching in below intense such one, whilst (Colmet- Vertisol and environments, soils humidity the similar (Chadwick which the and in retained above sites exhaustion of mm wet base composition two 1400 and the of The desilication have between threshold watersheds. and differs leaching MAP to processes distinctly the intense soil solutes thus with and affected 1965) of greatly Lagache precipitation Vertisol export has & and Abundant and regime Daage Nitisol 1965). water weathering the Lagache The to mineral March. contrast & enhanced in in (Colmet-Daage variation occurs little value composition very maximum shows mineral a 1b) which (Fig. in fieldwork to sites, prior season ‘dry’ the effie ocnrto a eemndo rne efsmlsacrigt h eegn brmethod fiber detergent the to according samples leaf grinded on determined was concentration fiber and combustion Leaf dry flash by measured analysis was Fiber leaves 103 in (DW; HNO concentration weight 10% dry Carbon in as ICP-AES. bead expressed by fusion & measured Nakagawa the Tombeur, were of Corn´elis, de dissolution K Nakamura, 1000 the 1992; at After Sanzolone calcinated 2020). & were Kitajima ashes (Chao of Li-metaborate mg and aeolian 100 Li-tetraborate from Then, particles calcination. potential the 65 remove after at to oven 450 an order in at in days four calcination ethanol for 70% dried were with They washed deposits. thoroughly were samples Leaf 2. 1. c i a g n concentrations C and K Mg, Ca, Si, analyses Plant kg -1 oetmt olwahrn tg Hriln1986). (Herbillon stage weathering soil estimate to ) acau officinarum Saccharum ° tal. et o 4hus h s ocnrto a eemndb egtdffrnebfr and before difference weight by determined was concentration ash The hours. 24 for C tal. et 3 01.Tpoladfla ape eecletdi rpiae nec ie For site. each in triplicates in collected were samples foliar and Topsoil 2011). , 03.Si rcse aeide e oteacmlto fscnayminerals secondary of accumulation the to led indeed have processes Soil 2003). lmn ocnrtoswr esrdb C-E.Tettlrsrei bases in reserve total The ICP-AES. by measured were concentrations element , -1 (Keeping ) ° samdlS–cuuao,btla icnetainvre eylittle very varies concentration Si leaf but Si–accumulator, model a is o or)adahfe r egtpretgsrsetvl (AFDW). respectively percentages weight dry ash-free and hours) 4 for C tal., et ° o 4hus olwdb uina 1000 at fusion a by followed hours, 24 for C 03 epn ee,2006). Meyer, & Keeping 2013; 3 ° n rne.S ocnrto a eemndafter determined was concentration Si grinded. and C 2 .1Mslto o h fe etiuain the centrifugation, After 6h. for solution M 0.01 2 etatbeS (CaCl Si -extractable ° 3 o i nagaht rcbewith crucible graphite a in min 5 for C h ocnrtoso i a gand Mg Ca, Si, of concentrations the , 2 n C ihaso- a with M 1 KCl and O 2 S)i osdrdto considered is -Si) ° o i na in min 5 for C Posted on Authorea 25 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158515652.24058515 — This a preprint and has not been peer reviewed. Data may be preliminary. etsl aestrto ntpol a 2 Adsl n 1 Ntsl ntewtetsts u reached but sites, wettest the in (Nitisol) 31% and (Andosol) Andosol) and 12% (Nitisol was 4.8 topsoils water, (cmol in in (Vertisol) CEC saturation 7.2 KCl. and Base Andosol) Vertisol. in (Nitisol, (Vertisol) 5.8 was 5.9 pH and soil 2a, Table in shown plants As in concentrations mineral and of properties level Soil 95% the at tests. different chi-square significantly Pearson’s Results were with letters tested MiniTab various were correlations and software Potential Fisher) the confidence. (LSD using differences significant performed least were analyses Statistical mappings elemental Si on signal) (Si analyses v2.10.8. pixels Statistical yellow mm GIMP of software 0.41 percentage the i.e. area with the pixels, squares, measured (200*200 five was squares same five the In on mm images. magnifications, per the three phytoliths the the dumbbell-shaped Bruker K of for a and Si side images with Stomata the abaxial fitted the on x150). on and acquired and obtained voltage mappings was x75 accelerating distribution Si kV Si elemental 30 The by microanalysis. at leaves backscattered-electron elemental working for the XL-30 detector with X-ray ESEM-FEG imaged eV paint were FEI mbar). silver 129 They with a (1.3 directly evaporator. bridged in mode MED010 and and detector low-vacuum tape Balzers (BSE) in carbon a tape double-side in and using carbon carbon-coated voltage slides to glass accelerating double-sided before on kV mounted with 30 were covered samples at leaf 600 slides Besides, Quanta glass ESEM on FEI 50 a spread at in were dried observed oven deposits was silica residue The Extracted times. microanalysis 3 X-ray repeated and centrifugation was for observation Rinsing tubes SEM min. polypropylene into 5 transferred for and rpm water 3700 deionized HNO at with 65% rinsed mixture 80 carefully the at was ultrapure baker residue Then, the tissues. organic in of added Fraysse portion gradually 1990; major 80 Kelly, the remove observation from at to microscopical (adapted order HCl and digestion 10% extraction wet with physical by baker for out used carried Corbineau was was 2009; site extraction , each The from deposits. samples silica leaf of three the of One deposits silica of kg extraction (g Physical percentages weight dry ash-free and hours) was fraction concentrations ADL lignin al. (Godin The and respectively fraction. hemicellulose ADL cellulose, ADLom, kg the (g The and percentages obtain ash). weight to residual dry h without as 3 ADL expressed for and % (ADF 72 90 subtraction acid by at sulfuric min (4): 550 15 fraction, at (ADF ADF for incinerated fibers the 7 detergent get pH the to acid at using h of determined buffer were contents phosphate 100 lignin) the 90 at mmol/L containing detergent hand, (ADL at 0.1 lignin other (1) min detergent extractants: the acid 15 following and On for lignin) ash). and 7 residual cellulose pH containing without at (NDF difference buffered lignin) 550 100 by and phosphate at at hemicelluloses mmol/L incinerated detergent cellulose, 0.1 containing was neutral (1) (NDF Soest extractants: fibers Van two detergent (2): neutral using of determined content was the hand, Schoelynck one 1967; the Wine, & Soest (Van 21) ial,tetresrcua opnnswr xrse sdywih gkg (g weight dry as expressed were components structural three the Finally, (2011). , ° o ihu h diino oimslt,()VnSetai eegn t100 at detergent acid Soest Van (3) sulfite, sodium of addition the without h 1 for C tal. et ° o ,adtems osalwdu ocluaetepretg fAFmadADLom and ADFom of percentage the calculate to us allowed loss mass the and h, 3 for C 03.Tngaso ahdadgiddla aeilwr rnfre noaglass a into transferred were material leaf grinded and washed of grams Ten 2013). , ° o ,adtems osalwdu ocluaetepretg fNDFom of percentage the calculate to us allowed loss mass the and h, 3 for C ° odsov abntsi n.Utaue6%HNO 65% Ultrapure any. if carbonates dissolve to C tal. et ° sln sterato eto n h eiu eandclru.The colorful. remained residue the and on went reaction the as long as C c tal et 04 05.Mr eal ntemto a efudi Godin in found be can method the on details More 2015). 2014, ° kg -1 o ihteadto fsdu ufie h D fraction NDF The sulfite. sodium of addition the with h 1 for C -1 W eete siae sAFmALm NDFom–ADFom, ADFom–ADLom, as estimated then were DW) ,21;VnSet 93 Godin 1973; Soest, Van 2010; ., a 58i h iio,4. nteAdslad5. nthe in 59.9 and Andosol the in 47.6 Nitisol, the in 25.8 was ) -1 AFDW). 4 α eka .4kVa ieetmgictos(x38, magnifications different 3 at keV 1.74 at peak ² ® fla ufc eecutdo h BSE the on counted were surface leaf of 81 en eecmae ae on based compared were Means 18.1. tal. et ² ° 3 admypstoe on positioned randomly ) ,()VnSetneutral Soest Van (2) C, 04 05.Bifl,on Briefly, 2015). 2014, , a rdal de in added gradually was ° -1 uig48h. during C W 103 DW; 3 3%H /30% 2 ° ° O o 4 for C o 1 for C 2 tal. et was ° C, et Posted on Authorea 25 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158515652.24058515 — This a preprint and has not been peer reviewed. Data may be preliminary. 5.) R (cmol CaCl TRB Vertisol. (55.1). the in 85% 14.Tbe2 ute hw httecneto o-xhnebeM agl otiue otetotal kg the (g concentration to Si contributed leaf The largely Mg non-exchangeable plants in of concentrations of content Mineral content the respective that the shows in reserve. gives occluded further non-exchangeable bases, cation 2b non-exchangeable each Table (cmol of reserve reserve for (104). the This one represents minerals. sum total soil Their the 2b). (Table from base content non-exchangeable exchangeable the Subtracting el)dmntdi evsadmliellrdpst curdol agnly(i.5) ag multicellular Large i). 5f, 5g). (Fig. (Fig. marginally size site Vertisol only 200 the that occurred to evidence deposits up gives multicellular deposits magnification and leaves fixed in at Nitisol dominated taken sequence cells) 5 ( the deposits Fig. in in dumbbell-shaped pictures increased small-sized 3 deposits of silica sets 3 of rods. the of long non-characteristic comparison and and or The size shapes 350 tissues small different to their inner of of 4e). up in lumens because cell cells (Fig. deposits probably walls, observed silicified large seen cell Very not to silicified were long correspond of cells and shape. may case guard phytoliths silica structures the stomata to silicified dumbbell-shaped from is Other corresponded for (iv) Deposits This them case 4d), veins. of silicification. the (Fig. Most from partial is structures cells to This 4f). silicified rod-shaped silicified (Fig. cells. (ii) elongated short phytoliths epidermal 4a), (iii) dumbbell-shaped and in 4c), (Fig. (v) exclusively walls and occurring (Fig. e), cell cells deposits 4c, silicified short (i) (Fig. and lumens 4): 4b) cell (Fig. (Fig. silicification structures cells by different long showed affected deposits area silica surface Extracted total deposits the silica site, extracted these Andosol of Between Despite site, Structures the Nitisol Vertisol 3a-i). in order i). the (Fig. the from stomata site h, leaves in Andosol silicified and f, the increased Nitisol of In significantly 3e, the proportion from cells. those epidermal high (Fig. to elongated compared a site silicified thin strongly Vertisol of appeared 200 3. (5-10) and veins to rows Fig. flat Andosol (up several in veins the included mappings wider and at and Si rows only flattened on more silicified shown veins, fully as prominent Nitisol site appeared sequence the which the on of in depended 20-70 increased greatly veins about cells of in short silicification and the lower a compared long Indeed, by as in 3k). site evidenced Andosol silicification (Fig. as the Vertisol) cells The from (Nitisol, silica leaves ones in in higher other than significantly two important was the less density were their to and stomata 3g-i) in (Fig. deposits signal mm Silica yellow 3a-j). per longitudinal (Fig. (number in site density arranged the and cells forms short size, and morphology long The their of epidermal cells to d). silicified guard according c, (iii) silicified types 2b, (ii) and 2a), different 2a) (Fig. (Fig. three (Fig. veins cells into rows silica in classified in located arranged be phytoliths stomata can dumbbell-shaped surface deposits (i) silica location: 2, and Fig. in shown As deposits silica leaf (cmol of Localization contents their of ( sites sum Nitisol the and concerned, are cations (56) Nitisol major the as far < noo (59) Andosol μ iewr omdb o3rw fsotbodeiemlcls(i.3-) h lumens the 3b-d), (Fig. cells epidermal broad short of rows 3 to 2 by formed were wide m > c kg 0) u afo h etsloe(44%). one Vertisol the from Ca but 50%), μ eeosre o h noo ie(i.5) n vnlre hn200 than larger even and 5h), (Fig. site Andosol the for observed were m -1 olwdtesm rn Tbe2) iio (35) Nitisol 2b): (Table trend same the followed ) 2 S m kg (mg -Si < etsl(9.Tedmnn aini uacn evswsKfo h Andosol the from K was leaves sugarcane in cation dominant The (79). Vertisol -1 Tbe3 olwdteodrNtsl(7) Nitisol order the followed 3) (Table ) c kg -1 -1 nrae ntesqec:Ntsl(27) Nitisol sequence: the in increases ) < μ olwdtesqec iio (16.8) Nitisol sequence the followed ) ogad150 and long m 50 < μ )adsotrdsiue ms rbbyfo tmt guard stomata from probably (most rod-spicules short and m) Andosol ² ftedmbl-hpdpyoih eecntn whatever constant were phytoliths dumbbell-shaped the of ) < 5 < Andosol etsl(i.3l). (Fig. Vertisol μ ieadwt eea el ntikeswere thickness in cells several with and wide m μ )wr oae ewe ogtdnlstomata longitudinal between located were m) < etsl(i.5-) o h iio site, Nitisol the For 5a-i). (Fig. Vertisol < Andosol c < < kg noo (14) Andosol noo (110) Andosol -1 olwdtesm sequence same the followed ) < < < noo (31.3) Andosol etsl rmnn veins Prominent Vertisol. etsl(63) Vertisol < < etsl(1.As (21). Vertisol etsl(114). Vertisol < μ < o the for m Andosol Vertisol Posted on Authorea 25 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158515652.24058515 — This a preprint and has not been peer reviewed. Data may be preliminary. nuneo efeiemlslcfiainta ietycnrl oeo h ao irltdfntosas functions Si-related major the of direct some a controls have directly conditions environmental that that silicification suggest epidermal we Therefore, leaf transpiration. McLarnon on Fitt, plant (Hartley, influence soil and in plant-availability 2015) Si cell through Wade during process & this saturation impact silica (Alexandre also In development on climate) stage (soil, 2018). depending leaf conditions Alexandre Elbaum process the regulated on & passive 2017b; depends a Kumar physiologically Elbaum cells is 2017a; short & and cells Elbaum Soukup active short & (Kumar, and (Motomura, Elbaum an long dehydration rate consistent Brami, is in transpiration is Milstein, of deposits cells This Kumar, independently silica veins. 2006; development, silica contrast, Vertisol leaf silicified 2004, in in of of Suzuki stage deposits 38.7% & one first Fujii the silica and the sites to during Andosol that three compared occurring showing in small the process mm 23.1% data is between per structures similar to literature stomata silicified very Nitisol with of these are number in by phytoliths higher CaCl affected % dumbbell-shaped significant in area 13.5 of increase a the number Although from with 3j). the contents, increases leaves.(Fig. opposite, largely Si sugarcane the deposits leaf in silica On by patterns resulting sites. affected biosilicification and area affect relative soil conditions the in climate 3l, Fig. and in soil show that As leaves highlight sugarcane soil also in the data patterns Our both silicification by cultivar on affected conditions negligible is environmental a of species case, Effect Hypothesizing plant any 7c). given In a (Fig. 1989) 1965). in content Kittrick Lagache accumulation Si & al. & Si leaf (Rai (Henriet that (Colmet-Daage controls solution stage corroborate season here soil weathering thus as dry soil in data well prolonged activity in our as a silica effect, Si 2) of high of Fig. occurrence bioavailability by 1, the controlled Table the (ETP, by is transpiration which promoted plant soil, as (Henriet and in availability banana stability soil water for smectite in accumulation affects earlier Si Si climate reported controls bioavailable important: as primarily by less stage conditions controlled far, weathering (Klotzb by climate is soil is, rice similar it Nitisol), which and in (Andosol, where 7b), sites site, cropped (Fig. wet Vertisol sugarcane content the the Si in 7a), in in leaf Thus, (Fig. the sampled concentration 7c). way leaves Mg same (Fig. sugarcane leaf the in the soil in impacts largest the affect directly (Table discriminate the not it to silicate Na-plagioclases does Despite indicator content primary and pertinent Mg soils. in (K) most non-exchangeable three poor feldspars the these very here of between is is stages disappearance Vertisol content weathering the near Mg line, non-exchangeable the Consequently, this by In 2). confirmed octahedral the Ca, 1965). whereas as in exchangeable Lagache Ca, minerals, occluded respectively. provides & total is weatherable carbonate (63), (Colmet-Daage Mg Vertisol of calcium non-exchangeable smectite the soil, 74% while of this complex and for exchange sheet In (27) accounts the Nitisol Mg. saturate which total to the Ca, contributes of in exchangeable which 16% than includes only higher represents massively times Mg Ca 1.6 exchangeable total and Vertisol, 3.8 the is In it (104): (cmol Andosol the reserve concentration non-exchangeable Si soil leaf The on climate and of soil f). concentrations of 6e, the Control (Fig. correction, concentration ash Si After leaf to 6d). Discussion correlated (Fig. not were concentration lignin Si and with hemicelluloses correlated negatively was and orcin ocnrtoswr iia o h he ieetsts(i.6) u ells concentrations ash cellulose After kg but g c). 6b), 6a, (455 (Fig. site sites (Fig. Vertisol different concentration three the Si the for with for lower kg correlated similar significantly (g were negatively were concentrations was They C concentration correction, cellulose (465). and Andosol kg C and (g Leaf (469) concentrations Nitisol Carbon sites wet 4). (Table (89) s ocnrtos( kg (g concentrations Ash concentration lignin and hemicellulose cellulose, Carbon, 05 ere,Dae piz wne evu 06 Issaharou-Matchi 2006; Delvaux & Swennen Oppitz, Draye, Henriet, 2005; -1 FW infiatydffrdi h re iio (374) Nitisol order the in differed significantly AFDW) ucher ¨ tal. et -1 tal. et W infiatyicesdi h re iio (54) Nitisol order the in increased significantly DW) 05.I otat nteVrio ie h mato olwahrn tg is stage weathering soil of impact the site, Vertisol the in contrast, In 2015). 08 ;Klotzb b; 2008a c kg -1 fwahrbemnrl Tbe2 s yfr h ags in largest the far, by is, 2) (Table minerals weatherable of ) -1 ucher ¨ W i o infiatydffrbtenlae rmthe from leaves between differ significantly not did DW) 6 tal. et tal. et tal. et 05,adpattasiainflxs(Euliss fluxes transpiration plant and 2015), > 09.Wieteslcfiaino ogand long of silicification the While 2019). 09,orrslsso htenvironmental that show results our 2019), noo (365) Andosol ² a bevdfrteAdslst,the site, Andosol the for observed was tal. et < > noo (66) Andosol etsl(5)(al 4) (Table (356) Vertisol 2016). tal. et < 08 b) 2008a Vertisol 2 -1 -Si et ). Posted on Authorea 25 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158515652.24058515 — This a preprint and has not been peer reviewed. Data may be preliminary. e otsi h pria fsi upesvns.Ide,sett ssal nsi tp rudo above or around pH at soil in stable is open smectite may Indeed, of here suppressiveness. impact reported soil The patterns H of biosilicification 2006). and appraisal and Steinberg neutrality plant the & the pH in in Olivain especially accumulation between routes Alabouvette, soil, Si new the on link 1999; of conditions (Alabouvette characteristics casual climate suppressiveness minerals abiotic and Soil The clay on soil far. of depending 1986). so interactions nature biotic established (Stotzky the soil on poorly and based property using however, be was, and specific to suppressiveness 1999) proposed soil this was (Alabouvette and suppressiveness predict smectite soil of to sensitive soilborne of occurrence by the concept composition induced of diseases the mineral plant eradication to suppressing developed the clay leading factor weakly to a 1986), as disease led (Stotzky smectite it this of pathogens whereas presence that the (smectite) showed highlighting clays crops authors swelling various These in rich (1963). cultivar soils The Martin, banana on 2016). & Delvaux cropped & Stotzky plants (Cornelis by in processes reported and by as (Fauteux type caused soil type e.g. disease, on (see Panama depends diseases of largely spread fungal soil in a.o. bioavailability against Si context, protection plant al. enhance plant et in functions Silicon-induced suppressiveness? soil on insights modulate the New (Schoelynck may Therefore, the earlier concentration supporting 4e). reported veins, Si (Fig. reinforcement silica leaf forming study and on those Klotzb this beneath biopolymers transpiration probably in most just C-rich plant cells, soils layer between and epidermal Si-rich balance soil cell (Yamamoto in from in cellulose the epidermis plants bioavailability of in adaxial synthesis Si in the located the the deposits is both silica in Si of with cells control in concerned short (Guerriero deprived cell in deprivation plants the whose Si and to biopolymer in for C-rich epidermis synthesis made kg role the abaxial are cellulose (g compensatory is ash-corrections in the concentration cellulose a no increase that deposits, cellulose when The supports silica leaf Si 2016). This decreasing by the 6d). with C (Fig. However, increases of concentration synthesis 2012). Si dilution Leishman leaf the components increasing highlighting & structural with sites, (Cooke of 3 8) types the of two (Fig. biosynthesis between the these differ than between not energy-consuming occur does less may is tradeoffs silica (Schoelynck 1983), of (Raven deposition the biopolymers (Kitajima Since leaves C-rich plant 2016). in Westbrook strength biomechanical & the Wright for mechanical responsible majorly leaf components is structural of Cellulose as analysis This cellulose complete plates. and rigid a silica local between with the Balance form conditions, of to controlled tissues stomata under plant cell and deep several tested involve of cells properties. be also thus deposits silica now but compact are cells Moreover, should form in and epidermal surfaces. could hypothesis deposits thus concern tissues, leaf that along only to leaf veins all not compared deeper reinforcement running columns longitudinal fertilization. large also columns role in Si silicified of but mechanical concentrated rigid under formation cells especially continuous a erectness are the epidermal play leaf they isolated that Indeed, in to only epidermis. hypothesize increase prone no we formation the more concern the Here, explain promote probably larger, to cells. site much crucial silicified Vertisol are be joint the They could in concentrations more deposits Si transpiration with leaf plant silica increasing deposits, and multicellular with silica soil increases leaves in larger bioavailability the of from Si (Yamamoto extracted Larger fertilization deposits 5). Si silica after (Fig. of improved size was the leaves that rice show of erectness leaf al. the et that demonstrated been has (Meunier It stress UV and water or (Cai 2009) intrusion pathogen against defense ucin,tehg iaalblt fS nsi ih hscnrbt otesprsieeso ihbase high of suppressiveness the to contribute thus might soil in Si of bioavailability high the functions, 02 Kido 2012; 05 atu,Can ezl,Mnis&Beagr20) nteohrhn,i ie climatic given a in hand, other the On B´elanger 2006). & Menzies Belzile, Chain, Fauteux, 2005; ucher ¨ tal. et tal. et rsMichel Gros tal. et 4 SiO 2018). 00 Klotzb 2010; 4 05,tepoesbigsildbtd(Bauer debated still being process the 2015), ocnrto vr1m nsi ouin eas ftepatpoetv Si-induced protective plant the of Because solution. soil in mM 1 over concentration nalohrsis hsnvlywsfloe olwd yanme fsuison studies of number a by worldwide followed was novelty This soils. other all in ucher ¨ uaimoyprmcubense oxysporum Fusarium tal. et tal. et tal. et 08 Schaller 2018; 07 Coskun 2017; 08,hrioy(pti 09 epn,Keaa Bruton & Kvedaras Keeping, 2009; (Epstein herbivory 2008), 7 tal. et tal. et 2019). 09.Hr,teahfe concentration C ash-free the Here, 2019). tal. et ae4o aaa a oendb soil by governed was banana, on 4 race 01 Cooke 2011; tal. et tal. et 00 Suzuki 2010; tal. et -1 02,corresponding 2012), tal. et FW decreases AFDW) 06.Orresults Our 2016). 02 Kitajima, 2012; tal. et 2012; tal. et Posted on Authorea 25 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158515652.24058515 — This a preprint and has not been peer reviewed. Data may be preliminary. rw nte3dffrn ol.Bt atcesz n ubro ag atce nraeovosyfo the 200 from than obviously larger increase structures particles indicate large (i) and of number (f) images and in size arrows particle The Both Vertisol. the soils. to different in Nitisol 3 arrows the The in structure. grown this on length 5: in The Figure attached structure. phytoliths. lumens rod-shaped dumbbell-shaped cell 350 silicified indicate silicified to a 35 (f) up indicates around structure image respectively (d) multicellular shows indicate large image arrow (c) a vertical in image indicate the arrow (e) in The image arrows lumens. in vertical the arrow cell and horizontal for horizontal and (%) The cells of cells. pixels short number long yellow silicified the silicified of counting indicates area for (b) Relative used image area 2. the 4: MAG visualizes and Figure mm (a) 1 in per pixels. MAG square phytoliths yellow red for and (DS) The soils stomata dumbbell-shaped (l). phytoliths, three of magnifications the and Number for soils (a-i). three (k) magnifications stomata direct cells and three long (j) at of sites, made three veins the indicate (c) 3: and (b) Figure signals in (d) arrows yellow respectively. vertical and cells, and (a-c) short horizontal BSE and The white stomata. Intense Leaf (d). and (b). Si phytoliths in of deposits fields mapping silica elemental sugarcane EDX indicate b). selected with in image the bars combined in (green and 2017 2017 May April 2: 13th to Figure to January 11th from from measured place as took ETP sampling Daily (a). sites manuscript. the wrote 1: authors Figure FdT, the fieldwork. All conducted data. BD analysed and legends CVL FdT Figure FdT, experiments. research. performed the PC designed and and BG planned CVL, CVL and for BD (ULiege) J-TC, interest. (CAREM) FdT, of Microscopy conflicts in declare authors Education data. the contribution and meteorological of Authors Research to None Applied access microscopy. for for electron France Neuchˆateau, Agneessens to Center Meteo Richard of access the laboratory, and Durviaux providing station soil laboratory, to Alexis UCLouvain research CRA-W grateful and the the CIRAD are in Hardy in We the help Brieuc analysis her thank of biochemical for We staff the Iserentant work. Anne for the laboratory assistance, and and field field Dorel their in for Marc assistance Dr. their for to Guadeloupe, gratitude our express We link identical sugarcane mechanistic play Acknowledgements in not casual do concentration (Soukup any constituents cellulose siliceous plants on and in in conclude lignocellulosic decrease roles since cannot investigated the physiological further we and be Yet, magnitude should Si-related leaves. This biosilicification leaves. and of sugarcane in leaves leaves in increase plant in biosilicification the cellulose of between on deposits of reinforcement mechanical synthesis silica role lower the of crucial on a size a effects play and resulting thus (Coskun localization their functions factors the on in environmental likely affect accumulation These and climate Si accumulator. patterns impact and high-Si strongly soil transpiration a that plant sugarcane, show and stage further weathering We soil plant. that corroborate data Our (Stotzky investigation. ecology microbial further however, on requires, properties Conclusion smectite hypothesis of challenging effects This the 1999). to Alabouvette addition 1986; in soils, clayey swelling saturated otl rcptto n eprtr saeae vr2 er fo 93t 08 ntethree the in 2018) to 1993 (from years 25 over averaged as temperature and precipitation Monthly S-VSMiae efre nasgraela aailsrae rmteVrio ie(a-c) site Vertisol the from surface) (abaxial leaf sugarcane a on performed images BSE-LV-SEM anfiainsre fBEL-E iw fslc tutrsetatdfo evso sugarcane of leaves from extracted structures silica of views BSE-LV-SEM of series Magnification obndBEL-E/D-impigiae faailsrae fsgraelae from leaves sugarcane of surfaces abaxial of images mapping BSE-LV-SEM/EDX-Si Combined S-VSMiae fetatdslc oisfo etslsgraelae.Tearwin arrow The leaves. sugarcane Vertisol from bodies silica extracted of images BSE-LV-SEM tal. et . 09.W ute ihih htteices nla icnetaincreae with correlates concentration Si leaf in increase the that highlight further We 2019). h oiotladvria rosi a niae epciey dumbbell-shaped respectively, indicate, (a) in arrows vertical and horizontal The tal. et 2017). 8 μ ogad150 and long m μ .Tearrows The m. μ ieand wide m ² Posted on Authorea 25 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158515652.24058515 — This a preprint and has not been peer reviewed. Data may be preliminary. al 1 Table Hasan from straw rice are Tables others the All study. orange. this in of 2001 Shen leaves yellow, sugarcane in the 1993 are dots Blue (n=103). olp pH pH Soil bases non-exchangeable of contents (TRB), Bases bases. non-exchangeable in of Reserve reserve CaCl Total total (BS), content), Na), saturation exchangeable K, base – Mg, (CEC), (total capacity (Ca, exchange contents cation elemental (SEB), (CaCl bases Si exchangeable extractable of sum Na), K, 2 Table (1965) Lagache 5 4 3 2 1 Vertisol Andosol Nitisol Group Soil Reference 8: Figure (b) brown). (a) (dark Vertisol 7: and Figure (orange-brown) Andosol brown), (light Nitisol the kg g and indicate cells. (h) short image 6: in and Figure arrows cells diagonal long and horizontal lumens, vertical, cells The attached structures. respectively multicellular show (g) image in acltdfo otl n eae(0dy aa(06 2017) (2016, data (10-day) decade and monthly from calculated 2017) (2016, Temperature Annual Mean Precipitation Annual Mean 2014) (IUSS key WRB ulfidfloigUD’ olSre aoaoySa 21) ae ndt rmCle-ag & Colmet-Daage from data on based (2017), Staff Laboratory Survey Soil USDA’s following qualified uacn efS ocnrto spotdaantsi CaCl soil against plotted as concentration Si leaf Sugarcane 1 -1 eetdsi properties: soil Selected : oegnrlcaatrsiso h he ie n uacn cultivars. sugarcane and sites three the of characteristics general Some : FW(,d ,f gis icnetain(ngkg g (in concentration Si against f) e, d, (b, AFDW lto efahcnetain( kg (g concentration ash leaf of Plot H lt ftecro a ) ells c ) inn()adhmcluoe()i kg g in (f) hemicellulose and (e) lignin d), (c, cellulose b), (a, carbon the of Plots 2 C aM acmol Na K Mg Ca KCl O 61 16 61 16 61 16 Coordinates lto uacn efM ocnrto gis h otn fnnecagal gi soil. in Mg non-exchangeable of content the against concentration Mg leaf sugarcane of Plot ° ° ° ° ° ° 27’29”W 25’32”N 35’33”W 02’59”N 38’04”W 10’36”N 2 tal. et S) o CaCl For -Si). 98i re,Abou-El-Enin green, in 1998 , (asl) m Altitude 8limestone andesitic 18 andesitic 150 163 kg (cmol bases Exchangeable (a) -1 2 S,S r ie ne brackets. under given are SE –Si, ) vrg n3 auso H otnso xhnebebss(a Mg, (Ca, bases exchangeable of contents pH, of values (n=3) Average c rock parent Soil (Pleistocene) (Eocene) ash (Pliocene) ash kg (cmol bases Exchangeable -1 gis efS ocnrto gkg (g concentration Si leaf against ) -1 ) c 9 kg (cmol bases Exchangeable mMAT mm MAP 222. 20utcB80689 ustic R579 R570 perudic 42.0 udic 34.2 26.7 34.5 25.3 1272 25.4 3170 2910 tal. et -1 ) c -1 2 2 W nlae rmsgrae rpe on cropped sugarcanes from leaves in DW) 99i e n Agbagla-Dohnani and red in 1999 , xrcal icnet(CaCl content Si extractable kg (cmol bases Exchangeable -1 SEB ) (b) c vrg n3 auso total of values (n=3) Average 3 kg ° C -1 -1 1 c nthe in ) mm/decade ETP E BS CEC kg cmol Poaceae 4 -1 2 -1 -Si). c W(,c) (a, DW tal. et tal. et family regime moisture Soil mg % 2 2 – , , 5 Si CaCl kg

-1 Cultivar

2 - mg kg -1 Posted on Authorea 25 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158515652.24058515 — This a preprint and has not been peer reviewed. Data may be preliminary. etsl5. 9642361391. 17382863 27 104 2.8 1.9 5.3 3.8 4.4 3.7 41.7 13.7 81.5 14.7 13.7 7.2 113.9 110.0 35.4 3.6 0.76 1.70 5.5 2.0 60 51 cmol 17 31 K 23 18 4.2 kg (g percentages Mg 5.0 weight dry 6.2 56 as 59 kg 12.5 expressed (g samples, percentages leaf weight (0.1)a (0.4)b sugarcane 13.2 11.8 the (0.2)b of 1.5 trations (0.0)b 4 (0.1)a Table 1.2 2.2 Ca (0.3)a 7.0 (0.1)b (0.1)b (1.1)a 2.6 2.1 21.0 (0.2)b 14.7 (0.3)c 7.0 Vertisol Si Andosol Nitisol Site are SE ratio. atomic 49.6 Mg/Ca and Sc, 83.6 in proportions 15.6 cationic 55.1 (Sc), contents. K mineral Mg, leaf the Ca, 31.3 for cations brackets of under sum given K; 16.8 Mg, Ca, 86 3 Table 12 59.9 31 5 47.6 cation. 4 51.2 25.8 3 5.7 (b) 0.8 8.1 56.6 0.2 15.8 Vertisol 11.5 0.4 0.1 Andosol (cmol concentration elemental Total Nitisol 1.3 7.9 1.8 Soil 2.1 42.1 1.9 2 2.1 1 5.9 (a) 4.3 4.8 7.2 4.8 pH Vertisol 5.8 Andosol 5.8 pH Nitisol Soil h o-xhnebecto otn stedffrnebtentettladecagal otn o each for content exchangeable and total the between difference the is content cation non-exchangeable The u ftecnet fnnecagal ae TB–SEB). – (TRB bases non-exchangeable of contents the of 1986) Sum (Herbillon cations alkaline-earth and alkaline major of contents total the of sum the is TRB BS=SEB/CEC*100 bases exchangeable of Sum vrg aus(=,S nobakt)o s,cro,cluoe eiells n innconcen- lignin and hemicellulose cellulose, carbon, ash, of brackets) into SE (n=3, values Average : iea otnsadblne nsgraelae:aeae(=)vle ffla otnso Si, of contents foliar of values (n=3) average leaves: sugarcane in balances and contents Mineral : aM acmol Na K Mg Ca kg (g content mineral Leaf -1 ) -1 AFDW). kg (g content mineral Leaf -1 ) kg (cmol bases Exchangeable -1 ) c kg (g content mineral Leaf -1 ) kg (cmol bases Exchangeable c -1 kg ) c -1 10 oa lmna ocnrto (cmol concentration elemental Total ) kg (g content mineral Leaf (0.3)ab -1 kg (cmol bases Exchangeable Sc ) -1 ) c kg 94 54 0.35 40 15 44 79 kg (cmol bases Exchangeable -1 -1 SEB ) c c -1 nS (%) Sc in proportion Cationic aM K Mg Ca W n/rahfe dry ash-free and/or DW) 1 E BS CEC c kg nS (%) Sc in proportion Cationic -1 oa lmna ocnrto (cmol concentration elemental Total ) 2 nS (%) Sc in proportion Cationic Si CaCl (11.0)a (1.6)b (0.8)b

2

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Mg/Ca Ratio 55.1 31.3 16.8 (11.0)a (1.6)b (0.8)b c kg -1 oa lmna ocnrto (cmol concentration elemental Total ) c kg -1 TRB ) c 3 kg -1 o-xhnebebss(cmol bases Non-exchangeable aM acmol Na K Mg Ca c kg -1 ) 4 o-xhnebebss(cmol bases Non-exchangeable c kg -1 ) 4 o-xhnebebss(cmol bases Non-exchangeable c kg -1 ) 4 o-xhnebebss(cmol bases Non-exchangeable c kg -1 ) 4 o-xh reserve Non-exch. c kg -1 5 Posted on Authorea 25 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158515652.24058515 — This a preprint and has not been peer reviewed. Data may be preliminary. hoTT azln ..(92 eopsto techniques. Decomposition (1992) R.F. Sanzolone 65–106. soils. & D.M. volcanic Hendricks T.T. of & functioning Chao W. biogeochemical Elliott the Crawford on C.G., climate Olson 195–223. of K., Ziegler impact E.F., The of Kelly (2003) mechanisms R.T., Gavenda cytological incidence O.A., rust and brown Chadwick decreases Physiological fertilisation Silicon (2008) (2013) X. A.R. Junior Zhu sugarcane. Gomes in & & L. J. Amorim de, Yang M.S. disease. Camargo R., blast Zeng against rice S., in Luo resistance D., silicon-induced volcanique Gao massif du K., 000 rice Cai 20 g´eologique Antilles). difference 1: Carte Petites (1987) of Guadeloupe, D. parts Westercamp Soufri`ere la (D´epartement & various la de M.P. de of Semet J., contents Dagain silica Transport, G., and Boudon plants: lignin land cellulose, cultivars. in the L.) mineralization sativa in (Oryza silicon Variation plants and (2001) rice P.-S. Calcium of Bonilla (2011) structure I.M. canopy Weiss and function. & Growth and R. (2002) structure T. application. Elbaum si P., Ajiki by & Bauer affected K. as Suzuki conditions H., field Fujii under grown K.I., leaf and Kakuda grass water of humidity. H., leaf Effects atmospheric Ando of (2019) continental J. signature of Roy isotope proxy oxygen . . new . triple 4613–4625. a C., the for Sonzogni on S., insights alternation Devidal phytoliths: light/dark C., Piel and A., development Landais anatomy, E., situation. Webb European A., the Alexandre diseases: plant Pathology of control Plant (2)b Biological of 50 (2006) (1)c C. (1)a Journal Steinberg European 43 56 & C. Olivain soils. C., disease-suppressive Alabouvette of example an (2)b soils: 46 Pathology suppressive (1)c (1)a Plant wilt and 40 53 Fusarium chemical (1999) C. degradability, (0.7)b Alabouvette 343 sacco straw. (0.7)a In rice 360 European (2001) (5)b of M. 341 varieties 15–27. 15 Doreau of (7)b A & composition 312 Cl´ement : G. morphological (0)a Lignin plants Nozi`ere P., Ecotoxicology (5)ab 337 in (2015) 322 A., toxicity M. Kashif Agbagla-Dohnani metal (0.1)c heavy 356 . of . . alleviation F., silicon-mediated Abbas (1)b M., of (4)a varieties.review. 365 Ibrahim HemicelluloseHemicelluloseLignin Mechanisms 374 rice M., Safety Zia-ur-rehman Environmental (2)c 53 M., and 324 Rizwan from S., Ali ammonia M., Cellulose (1)b Adrees anhydrous (4)a 341 354 without, digestion and fibre (1)a and 500 with, composition chemical Technology Cellulose straw in Differences (1)b rice (1999) 498 D.J. (1)c of MacKill 496 & J.G. Fadel (2)b O.H., 455 Abou-El-Enin Carbon (1)a 465 (1)a 469 References (3)a 89 Carbon (1)b 66 (1)c 54 Vertisol Andosol Ash Nitisol name Site ctxclg n niomna Safety Environmental and Ecotoxicology 79 WD FWD FWD FWD AFDW DW AFDW DW AFDW DW AFDW DW DW rpProtection Crop 129–136. , 28 57–64. , ln Science Plant h hlpiearclua scientist agricultural Philippine The 53 72–79. , 180 114 746–756. , 329–341. , 119 11 186–197. , hsooi Plantarum Physiologia olSineadPatNutrition Plant and Science Soil 84 ora fGohmclExploration Geochemical of Journal RM Paris BRGM, nmlFe cec n Technology and Science Feed Animal 126–137. , igocecsDiscussions Biogeosciences 134 . nmlFe cec and Science Feed Animal 324–333. , hmclGeology Chemical 48 Australasian 429–432. , 202 44 16 94 , , , , Posted on Authorea 25 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158515652.24058515 — This a preprint and has not been peer reviewed. Data may be preliminary. seseto h eegn bradteislbedeayfie methods. metrological fiber comparative dietary biomasses: insoluble plant the in Lignin and (2015) fiber J. detergent Delcarte the & of P. assessment Gerin R., Agneessens B., Godin correla- methods. biomasses: fiber plant dietary in and the carbohydrates and Structural Silicon 62 (2014) fiber J. detergent between Delcarte the Trade-Offs & between P. tions Gerin (2016) R., Agneessens S.N. B., Plants. Johnson Godin Phenolic-Rich plant & on Herbivores of P.G. Root Ecology reactivity of Allsopp Chemical Performance and of N., Enhanced Explain chemistry Sallam may Surface Defenses J.R., Phenolic (2009) Powell J.D. A., Meunier Frew & resistance solutions. J. aqueous disease Schott in plant O.S., phytoliths and Pokrovsky Silicon F., the Fraysse (2005) in B´elanger R.R. Si & of fungi. J.G. pathogenic role against Menzies protective R´emus-Borel W., The F., (2006) Fauteux B´elanger R.R. & J.G. Menzies 17559. F., pathosystem. Belzile mildew F., Arabidopsis-powdery plants. silica Chain in tissue F., silicification plant Fauteux biological of of use mechanism The possible (2005) A 1–7. A.P. (2015) Schwab C. & Exley M.K. Banks transpiration. K.C., estimating Benke for B.L., content plants. Dorsey in K.W., roles Euliss manifold Its Silicon: (2009) E. Epstein Silicon. (1999) E. biology. Epstein America plant of in States silicon United of the anomaly of plants. The in (1994) The influx E. silicon (2019) Epstein and B´elanger R.R. aquaporins . of . . evolution Ecology Molecular J.F., tional (2016) Ma B´elanger R.R. & O., R. Reynolds loop. biology. Deshmukh J.G., plant feedback in Menzies silicon role H., silicon’s biological Sonah of the controversies R., Deshmukh drive D., processes Coskun Soil (2016) B. Delvaux Ecology & J.-T. Cornelis analysis. meta- isotopic concen- carbon a phytolith for pure addition: 179–185. suitable producing Towards are , silicon (2013) that G.M. with plants Santos from & stress A. trates abiotic Alexandre P.E., of Reyerson R., alleviation Corbineau Consistent (2016) M.R. Leishman analysis. & from Evidence J. defences: Cooke carbon-based types. and soil silicon contrasting genes. foliar between of to Tradeoffs communities geoscience (2012) vegetation silicon: M.R. plant Leishman & of J. ecology Cooke functional The (2016) S.E. Hartley Ecology & Functional J.L. volca- DeGabriel d´eriv´es roches J., sols Cooke de de groupes quelques fran¸caises. Carat´eristiques antilles de (1965) aux P. soils. niques Lagache in & minerals F. of Colmet-Daage occurrence and Processes formation, and Alteration, Properties Sciences: (2012) D.. Soil Lowe & G.J. Churchman 5609–5616. , 30 ucinlEcology Functional 1298–1310. , 30 1277–1285. , 30 42 1270–1276. , 768–771. , ESMcoilg Letters Microbiology FEMS nu e.ln hso.PatMl Biol. Mol. Plant Physiol. Rev.Plant Annu. 30 air RTM S´erie P´edologie ORSTOM. Cahiers hmclGeology Chemical 1340–1357. , 91 11–17. , clgclEngineering Ecological 1 20–72. , rceig fteNtoa cdm fSciences of Academy National the of Proceedings e Phytologist New Oikos 258 12 249 naso ple Biology Applied of Annals 197–206. , 121 rceig fteNtoa cdm fSciences of Academy National the of Proceedings 1–6. , 25 2052–2060. , 221 343–348. , eiwo aaooayadPalynology and Palaeobotany of Review ora fArclua n odChemistry Food and Agricultural of Journal 3 67–85. , 91–121. , 50 641–664. , Cellulose rnir nPatScience Plant in Frontiers 155 22 155–160. , 2235–2340. , 103 adokof Handbook Functional 17554– , Journal Func- 197 6 , Posted on Authorea 25 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158515652.24058515 — This a preprint and has not been peer reviewed. Data may be preliminary. azO 21)Slcncneti ln ucinltat mlctosi hnigworld. changing a in implications trait: functional plant a is content Silicon 88–94. (2019) O. the in Katz activities enzyme stress. strengthens drought and photosynthesis under improves xanthoxylum 76–86. Silicon Zygophyllum (2016) xerophyte Animals. X. succulent Zhu and C3 & Plants, W. Zhao Soils, J., Kang In Silica (1967) K.A. Handreck 107–149. & L.H.P. Jones an 2014. classification along soil Rome for Nations, pedicellatum base Zirihi-Guede reference Pennisetum World & (2014) species IUSS C. grass Contoux the P., in Dussouillez Niger. variations 220 South M., silica in Saadou gradient biogenic J.D., evapotranspiration Intraspecific Meunier (2016) D., N. Barboni composition silicon I., the in Issaharou-Matchi variation Phylogenetic phases (2005) developmental M.R. Broadley the & plants. on A. of Mead study P.J., ultrastructural White M.J., An Hodson in of (1985) sequestration glumes D.W. the carbon Parry of in & silicification phytoliths A.G. and lumen Sangster and M.J., wall Hodson cell of importance hypothesis. Janeiro, relative A low de The soil: Rio of (2019) EMBRAPA, horizon M.J. I. diagnostic Hodson Part the Ultisols. in of present Utilization minerals and primary rization, weatherable weatherable of soils. of estimation clay reserve soils. activity Chemical The ash volcanic (2008b) (1986) B. in A. Delvaux silicon Herbillon & biogenic S. of Opfergelt accumulation silicon M., the of Dorel uptake impacts N., and silicates Jaeger distribution De Effects, C., (2006) conditions. (Musa Henriet B. controlled banana Delvaux under & in spp.) R. content (Musa Swennen silicon banana I., Guadeloupe. Leaf in Oppitz in X., (2008a) soils Draye ash B. C., volcanic Delvaux Henriet of & stage X. weathering Draye the reveals M., spp.) Dorel Mackay. at Bodarw´e L., trials C., silicate plant Henriet calcium Technologists. rice the Cane of of Sugar aspects resistance of Some Society lodging (1975) Queensland L. influencing Chapman components & Chemical M. Haymsom (1993) vitro. in I. digestibility Goto straw inter- & its and M. and intra- Shimojo surface: supply. S., acid leaf silicon and Hasan silicic the damage Defending does to response (2015) How in R.N. tough. grasses Wade in and Science & deposition Rough silicon E.L. in (2018) McLarnon differences C. specific R.N., Exley Fitt S.E., & Hartley K.L. infection? Moore Matrix. fungal I., Extracellular from Stokes Plant horsetail C., protect the Law and G., Silicon Guerriero (2016) S. Legay & Science J.-F. Plant Hausman T0792 G., (2011) J. Guerriero Delcarte & lignocellulosiques. 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( (g) (d) a) NIT IS O L 200 1000 500 µ m µ µ m m (h) ( (b) e) A 19 NDO S O L 200 1000 500 µ µ m µ m m ( ( (f) i) c) VERT IS O L 1000 200 500 µ µ m µ m m Posted on Authorea 25 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158515652.24058515 — This a preprint and has not been peer reviewed. Data may be preliminary. (a)

Leaf Mg (g kg−1) (e) (c) (a) -1 -1 -1 0 1 2 3 Lignin (g kg AFDW) Cellulose (g kg DW) Carbon (g kg DW) 310 330 350 370 390 450 470 490 510 30 40 50 60 Non e Nitisol 0 25 0 0 p<0.01 r =-0.970 p<0.01 r =-0.945 xc hang eab 5 Andosol 5 5 50 le Mg(cmol Si (gkg 10 10 10 75 V c kg er -1 tisol −1 D ) 15 15 15 W) (b) Leaf Si (g kg−1) 10 20 30 0 Nitisol 20 20 20 Non e Nitisol 25 xc 25 25 25 hang 20 Andosol (d) eab (f) Andosol (b) 50 -1 -1 -1

le Mg(cmol Hemicelluloses (g kg AFDW) Cellulose (g kg AFDW) Carbon (g kg AFDW) 450 470 490 510 310 330 350 370 390 310 330 350 370 390 75 0 0 0 p<0.01 r =-0.928 V c kg er Vertisol tisol −1 ) 5 5 5 (c) Leaf Si (g kg−1) 10 20 30 0 Si (gkg 0 10 10 10 Nitisol 20 -1 D CaCl 15 15 15 W) Andosol 2 -Si (mgkg 40 20 20 20 60 −1 ) V er 25 25 25 tisol 80 Posted on Authorea 25 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158515652.24058515 — This a preprint and has not been peer reviewed. Data may be preliminary. 21