Posted on Authorea 20 Apr 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158739590.04450109 | This a preprint and has not been peer reviewed. Data may be preliminary. rncito atr eesqetal erie uigse ln vlto ocnrldvretat including traits diverse control to detected evolution factors plant was transcriptions in seed characterized partitioning during polymorphism Well recruited carbon diversity. high sequentially balance were morphological a to factors observed (iii) transcription complexes level, the and regulatory to intra-species master diversity relation the in molecular natural in interact At the intra-species genes of few and the evolution functions. a inter- summarize the for ecological we observed (ii) identified diversity, Here, the myxodiaspory cell recently species. underlying whether secretory the several myxospermy unclear mucilage in in is and disappeared involved mucilage It it the actors fruit why stages. (i) the and seedling on or origins early advances (myxospermy) evolutionary seed recent the multiple the in or of roles plant imbibition one play the around has can upon present therefore adhesion. released often and and is protection features, mucilage (myxocarpy), including a diverse functions species, with numerous providing myxodiasporous trait environment, In plant the with widespread contact a in structures is mucilage polysaccharidic The Abstract [email protected]) Castanet- ([email protected]; 31326 authors Toulouse, corresponding * Universit´e de V´eg´etales, UPS, Sciences CNRS, France en Tolosan, Recherche de Laboratoire Dunand* Christophe Burlat*, particular Vincent S´ebastien for Viudes, functionS ecological versatile generate mechanisms molecular environmentS diverse evolution: recently support. mucilage establishment but Seed plant dissemination and seed control and micro-organism germination Rosids. soil to across as conserved related such be mostly uncovered to were were seem seed functions mucilage seeds during exosystemic seed in recruited some of functions sequentially to their functions were and complexes ecological myxospermy, factors regulatory the including transcription Historically, traits relation master These diverse in in control genes seeds. interact to few thaliana evolution factors a Arabidopsis plant evolution for transcriptions the in the detected characterized (iii) partitioning was Well (ii) and carbon diversity, polymorphism diversity diversity. balance high cell natural morphological a intra-species secretory level, observed and mucilage intra-species early the inter- several the and the observed to in At mucilage the in disappeared functions. the underlying roles it ecological myxospermy (i) why play identified in and on recently can involved origins advances therefore actors evolutionary recent molecular and multiple the the (myxocarpy), or of summarize one fruit we has the myxodiaspory Here, or whether (myxospermy) mucilage unclear species. seed a is the contact species, It in myxodiasporous of stages. structures In imbibition seedling plant adhesion. the around and present upon protection often released including features, is functions diverse numerous with providing trait environment, plant the widespread with a is mucilage polysaccharidic The Abstract 2020 28, April 1 S´ebastien Viudes environments particular for generate functions mechanisms ecological molecular versatile diverse evolution: mucilage Seed aoaor ercecee cecsvegetales sciences en recherche de Laboratoire 1 icn Burlat Vincent , 1 n hitpeDunand Christophe and , 1 1 rbdpi thaliana Arabidopsis ed.These seeds. Posted on Authorea 20 Apr 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158739590.04450109 | This a preprint and has not been peer reviewed. Data may be preliminary. ol aesatdt eascae ihmxsem.Hwvr ueosMCtobxgnsfr a it form when genes and toolbox toolbox, MSC display this numerous can of genes However, individual origin and the myxospermy myxospermy. (Golz to clarifying contributions with functions turn unequal pleiotropic associated has in between that be species, network relationships regulatory to other orthologous complex in started identify genes have should and could analyses establishment genes mucilage Computational toolbox seed in MSC 2015). implicated myxodiaspory actors Dunand of molecular field & the under-studied in the of primarily on characterization focus performed deep to was is a studies review far, this rare ( So of Such field aim evolution. research overall angiosperms. unclear single The a across is to ecology). spread restricted or well It are development integrative species supposedly, as and on and species was, descriptions, actors comparative it. few morphological molecular it a require extrude standardized involved concern will reason and the not questions precise which of do evolutionary to characterization for these function(s) as or and mucilage to mucilage seed answer origins have Full connecting evolutionary studies not species. multiple 2018), several do Burton or in & orders one (Phan lost plant tree has phylogenetic studied myxodiaspory angiosperm the the whether along within easily 2018). all species be Hoffmann described supplement can several & been dietary has Gaiani Berger, and myxodiaspory Soukoulis, as North, properties if industry, 2012; Even chemical 2012c; Amid food and & Huang and (Mirhosseini physical pharmaceutical & respectively specific in Baskin biopolymer to developed has Baskin, and documented were mucilage Yang, was applications As investment 2012; several Western metabolic extracted, 2014). modification in this physical Ralet direct (reviewed species, by & conditions dispersal studied Saez-Aguayo seed of local and protection array the seed large as of such the functions In ecological the numerous fulfill of 2007b). structural 2-3% North and for & accounts biochemical Poll it molecular, composition instance, the biochemical for and as, the development, 2018; explored in Burton polysaccharidic seed have & particular released during Phan reviews in their MSCs Recent mucilage, and of of angiosperm 2018). (MSCs) characterization dynamics scientific of Burton wall cells majority the & cell a secretory its of in Phan mucilage and interest of described of in reported was the description diversity (reviewed myxodiaspory drawn was The years, the mixtures has 50 presence with last functions 2011). mucilage the associated ecological Leubner-Metzger In Seed orders putative the century. & its a from Muller over 1). and coming for (Weitbrecht, (Figure community ability species letter release same myxocarpy several to Darwin’s the in called species Charles while morphology of is myxospermy, ability a layer called The in is cell 1). early Figure epidermis upon outermost 2001; mucilage coat epicarp (Ryding extrude seed pericarp to fruit the fruit ability properties from the the physical mucilage or designates of coat seed diversity of myxodiaspory seed impressive range term the an wide The from conferring a imbibition by stems 2020). such secreted and In Krause leaves, be functions & 2000). can roots, various Knox Garbary (Galloway, compositions fruits, for wall-like & seeds, organs cell Nickrent as with around such Duff, mucilage organs it plant (Renzaglia, of non-vascular extrude reproduction kinds several diverging which and plants, early growth flowering sp.) during in Anthoceros protection produced dehydration (e.g. be as hornworts to as found such are groups mucilage, called also Mucopolysaccharides, Introduction evolution gene toolbox head MSC Running regulator, master evolution MBW its myxodiaspory, myxocarpy, of myxospermy, part roles, and diversity some words: mucilage recently Keys seed functions. but support. ecological of dissemination establishment its understanding ecological plant seed as the and the well advanced and control Historically, as have micro-organism germination studies soil Rosids. to recent as across related such These conserved uncovered mostly were be were functions to mucilage exosystemic seem seed seeds of in functions functions their and myxospermy, edmclg,itrseisntrlvraiiy nr-pce aua aiblt,ecological variability, natural intra-species variability, natural inter-species mucilage, seed aitv vlto fmxsem o iesfideooia functions ecological diversified for myxospermy of evolution radiative A : oa en&Huh 09) edmclg a ersn infiatmtblccost metabolic significant a represent can mucilage Seed 2019a). Haughn & Dean Sola, ˇ tal. et .thaliana A. 08.Teeoe ti eydffiutt lal dniygnscommitted genes identify clearly to difficult very is it Therefore, 2018). osiuigtes-aldMCto o Facz aoh,Burlat Ranocha, (Francoz, box tool MSC so-called the constituting .thaliana A. rbdpi thaliana Arabidopsis 2 edms Mcut ae,Koebre,Marion- Kronenberger, Ralet, (Macquet, mass seed e.g. Gits&Nrh21;Golz 2017; North & (Griffiths eei hrceiain hsooyand physiology characterization, genetic .thaliana A. tal. et 2018; Posted on Authorea 20 Apr 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158739590.04450109 | This a preprint and has not been peer reviewed. Data may be preliminary. cusete iutnosyi l rbdpi Sso eunilyi daetflxMC Fgr 2). (Figure wall MSCs cell flax differential scaffold domains by molecular adjacent development wall polysaccharidic-proteinaceous in seed The primary MSC sequentially peculiar modifications. modes during of localized or extrusion earlier and rupture prepared MSCs mucilage deposition the carefully polysaccharide Arabidopsis seed Indeed, is all in explosion 2). organized in occur (Figure This simultaneously trait differences the either subtle in of However, 75-fold occurs diversity to extrusion the up respectively. mucilage to increase to 2014), contributing mass leads and Neffati this volume & Finally, mucilage mucilage Yang seed a nature 2). the induces hydrophilic (Figure result, This the wall a pastoris First, As cell absorption. primary seed. water species. MSC the allows outwards all the mucilage in breaking the seconds pressure constituting swelling within mixture polysaccharide occur events the Whitehouse sequential of & (Vaughan imbibition, genera seed 90 covering Upon morphological species in pioneer Brassicaceae the illustrated by 200 also exemplified on is as diversity conducted 2018), 1971). Burton morphological MSCs & MSC mature (Phan the of columella years visible the survey models, called becomes over two and studied wall MSCs these species the secondary various Beyond in the polarized deposited 2). sequentially volcano-shaped is (Figure mucilage multilayered extrusion a complex after and a flax, wall In primary 2). (Figure periclinal extrusion outer and the the patterns development, between organization MSC mucilage various the (Figure the During maturation in seed as the well of as end modes. development the at seed layer during dead dynamics Miart a 2001; become environ- Western to the development 2; to seed in extrusion during mucilage exemplified differentiated enabling As layer is 2A). cell epidermal 1, coat (Figures seed ment outermost the Gorb to & Ultrastructural correspond (Kreitschitz MSCs 2019). reported Ralet also or was & microscopy North seeds force Botran, atomic hydrated Williams Poulain, and 2018; 2017; whole microscopy Wu electron & on scanning Qiao by performed analysis (Zhao, mucilage directly a of is with antibodies obtained extraction (MSCs) is staining easy epitope cells characterization (Ben-Tov section Further wall secretory sections or 2001). mucilage cell (Western on red, blue study with ruthenium toluidine to immunofluorescence with as and such by seeds stain mucilage hydrated polychromatic of on and presence generic performed the staining attest pectin to mucilaginous method convenient A plant seed on myxodiaspory on impact of Yang insights its Diversity 2012; recent as (Western discuss well 2012 we in as part, on reviewing constraints, last focusing last biotic the developed the and In are since abiotic pigments level. obtained facing development protective inter-species functions and and ecological intra- lipids numerous the storage mucilage both as at such briefly diversity in traits pioneer identified we their seed initially the First, genes, other between underlying with the comparison mucilage. of integration by seed origin evolutionary illustrated of occurrence the mainly concerning functions the actors literature diversity ecological molecular (i) structural various the on thaliana MSC of the survey evolution A. and a (iii) the mucilage in and (ii) seed information trait myxodiaspory), of expose complex of levels widely this multiple more in for integrate (and species. involved nutriments to myxospermy the is of required on diversity review depending the and advantages this provide ecological of & additional (Baroux will objective bring protection The endosperm will radiation mucilage and the the impermeability (Song and and confer mucilage 2019), embryo will seed Grossniklaus pigments the the while development, in and embryo pigments accumulating proper flavonoid Lipids lipids, storage 2017). between in Indeed, partitioning carbon analyses. computational dictates from merely myxospermy, with Dn,Jn,Too,Sur anta21)ad55fl in 5.5-fold and 2012) Iannetta & Squire Toorop, Jeng, (Deng, oe n h mrigflxmdl ntescn at epeettecretsau fthe of status current the present we part, second the In model. flax emerging the and model tal. et 2020). tal. et tal. et 08.Plschrd hmclaayi a epromd aiiae ythe by facilitated performed, be can analysis chemical Polysaccharide 2018). 09.Teetomdlseisilsrt h iest nteMCcl wall cell MSC the in diversity the illustrate species model two These 2019). .thaliana A. .thaliana A. edmclg stapd ihn paetsub-layering, apparent no with trapped, is mucilage seed and 3 iu usitatissimum Linum .thaliana A. eohtndeserti Henophyton tal. et flx,ti pcfi ellayer cell specific this (flax), ed,argltr complex regulatory a seeds, 2012c). .thaliana A. Gri lAloui, El (Gorai, aslabursa- Capsella Their . tal. et Posted on Authorea 20 Apr 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158739590.04450109 | This a preprint and has not been peer reviewed. Data may be preliminary. noedrc cosrsosbeo edmclg ytei,asml n erto (Francoz secretion gene and the assembly synthesis, in mucilage position seed whose of genes responsible regulatory (Golz regulators actors characterized master puzzling direct less upstream encode still characterized and mucilage is well hereafter, seed network including discussed proper factors regulatory further a transcription be to are will participate genes that that these of toolbox majority MSC (Francoz Arabidopsis A growing in continuously release the and constitute production They far. Shimada 2018; so Yu & Ji Sola Chen, in Zhang, physiology (Li, characterized MSC al. been and (Francoz have functional mucilage allowed genes genes have seed additional approaches 54 in global reported involved more genes reviews with together numerous genetics of reverse downstream characterization and and forward regulators of upstream years between Twenty separated be may myx- genes box species actors tool inter MSC The and 3.1. intra- the underlying variability actors natural molecular ospermy light the shed of inter-species will Evolution and we intra-species part, following through understanding. evolution the diversity mucilage in morphological seed reason, exemplified for this this as studies underlying For comparative families myxodi- mechanisms within the 1971). even molecular of Whitehouse and distribution polysaccharidic on & families clear-cut mucilage (Vaughan angiosperm simple and Brassicaceae the no for mode, along is traits extrusion there However, myxodiasporous mucilage asporous/non structure, organization. structural MSC may and including peculiarity diversity, composition additional of (Song an levels embryo tegument Finally, the tiple pericarp-seed and inner coat 2017). seed the Kester the via & seed Al-Amery the in to Phillips, exist attached Hildebrand, indirectly as (Geneve, remains such contact mucilage species the the myxospermous extrusion, Indeed, with After environment. parallels differentiation the interesting similar facing show the cells usitatissimum integument MSCs to seed outer and leading the of convergences mucilage by types evolutionary ene not different of and of examples pericarp, MSCs clear fruit) into are (non-dehiscent These achene the 2001). named (Ryding layer, cell fruit as 2010). such outermost Western species & myxocarpous Haughn for (Arsovski, by Interestingly, characterization understanding observed allowed dynamics mucilage microstructure wall seed cell mucilage (Yu Studying for seed interactions model 2018). of specific Gorb diversity polysaccharide-polysaccharide & the of (Kreitschitz demonstrated microscopy has electron species scanning multiple among ison te pce uha o example, Kreitschitz for pectinata 2; as such (Figure (RGI) species respectively rhamnogalacturonan other xyloglucans/cellulose, I Miart bound and type 2017; arabinoxylans layer branched Gorb RGI, adherent & poorly in an in enriched between enriched layers separated both trasted Macquet layer, is 2; non-adherent diver- mucilage (Figure observable a released pectin the and the to seed contribute Arabidopsis, the directly In to mucilage seed myxospermy. (Miart of of opening composition MSC sity polysaccharidic proper and in structure role The a play to proposed are organization Aguayo the enabling ˇ 08 Voiniciuc 2018; tal. et eiaotruncatula Medicago tal. et Ketciz&Gr 07 or 2017) Gorb & (Kreitschitz 09;Yang 2019b; .thaliana A. uiaeacmltsi h ue eiapeiemlclsdrn hase development. seed chia during cells epidermal pericarp outer the in accumulates Mucilage . 03 Francoz 2013; tal. et tal. et tal. et rmr aldmi osnn trst eucvrd(Kunieda uncovered be to starts loosening domain wall primary 08 a Wijk van 2018; 09.Ti yeo edmclg aibelyrn n opsto loeit in exists also composition and layering variable mucilage seed of type This 2019). tal. et and tal. et 09 Wang 2019; 07;Poulain 2007a; .orbicularis M. eiimperfoliatum Lepidium tal. et 09.I flax In 2019). tal. et lnaoovata Plantago tal. et tal. et 07.Teeoe yoisoyi ri htecmassmul- encompasses that trait a is myxodiaspory Therefore, 2017). tal. et 05 n 2gns(hn&Bro 08.Snete,12 then, Since 2018). Burton & (Phan genes 82 and 2015) Salvia h elwl fteedsemfrsamclg e between gel mucilage a forms endosperm the of wall cell the , tal. et tal. et 08.Teohrdwsra ee ftetobxmostly toolbox the of genes downstream other The 2018). 08 uid,Hr-ihmr,Dmr agn2019; Haughn & Demura Hara-Nishimura, Kunieda, 2018; 09 Fabrissin 2019; , h S oyacaii opsto n t internal its and composition polysaccharidic MSC the 4 and 05 oncu,Yn,Shit ¨n sdl2015). Usadel G¨unl & Schmidt, Yang, Voiniciuc, 2015; 09.I a,temclg scmoe yfu con- four by composed is mucilage the flax, In 2019). Hag ag un a a 2015), Lan & Cao Yuan, Wang, (Huang, (Tucker Artemisia tal. et tal. et tal. et 08 aigse uiaea excellent an mucilage seed making 2018) pce,temclg setue ythe by extruded is mucilage the species, 07 Yu 2017; tal. et 09 rnigtels o9 genes 94 to list the bringing 2019) 2019). avahispanica Salvia tal. et tal. et 07.Tecompar- The 2017). .thaliana A. .thaliana. A. 08 Takenaka 2018; tal. et tal. et Neopallasia 03 Saez- 2013; ci)ach- (chia) 2015). Recent and L. et Posted on Authorea 20 Apr 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158739590.04450109 | This a preprint and has not been peer reviewed. Data may be preliminary. 09.Tecmiain fseicMBadbL rncito atr oehrwt TRANSPARENT with together factors transcription bHLH and MYB specific of proanthocya- and combinations (anthocyanidins The biosynthesis flavonoid 2019). the 2012), Dolan (Lloyd & nidins) (Jones hairs root and trichomes regula- In master conserved a involves variability complex myxospermy the tory inter-species study angiosperms. angiosperm to across toolbox. The allow genes will MSC 3.3. toolbox mostly network the MSC regulation pressure of the toolbox selective genes of the myxospermy part target of the of genes downstream level, origin upstream their evolutionary the intra-species altered of the but conservation at regulators this Hopefully, that master suggests upstream also the It conserved mucilage species. seed other of production in the in implicated in as characterized characterized one a another (Fabrissin being and content (Soto- mucilage toolbox pectin seed MSC modification to the or to pointing belong positions, synthesis to implicated mucilage less (Fabrissin seed of trait background of characterized polygenic high actors previously the above of direct significantly orthologs downstream peaks are are five that content, genes Cerda mucilage box in tool implicated In MSC loci phenotype. seven mucilage seed the observed of the explaining (SNPs) polymorphisms nucleotide on conducted (GWAS) usitatissimum studies L. association wide genome recently, More phenotypes mucilage their species. proposed and myxospermous functions habitats mucilage for population seed function natural the the of (Macquet between one established myxospermy is (Voiniciuc be run-off of can water by association loss dispersal clear the seed river, explain a to to close is habitat Sha (Francoz related Kunieda and in occurring diversity easier natural be the should Indeed, still al. scarcer. genes are et are changes toolbox and because time MSC variability, scale natural the usitatissimum evolutionary inter-species on the the on pressure and than recent rather selection synthesis relatively intra-species of mucilage the effects in considering involved the when directly understanding genes and tar- box Detecting pressure tool selection MSC strong downstream a few release reveals a variability on natural geted myxospermy intra-species The 3.2. hssrnl ugssta ontemMCtobxgnsudrosrn eeto rsue u also but (Saez-Aguayo independent pressure, seven area selection geographical the strong different by two exemplified undergo only as genes in trait toolbox found fast-evolving mutants MSC a is downstream myxospermy that (Saez-Aguayo that populations suggests natural strongly studied 300 This nearly the considering and since synthesis and mucilage of namely (Saez-Aguayo box, lack MODIFIED2/BETA-GALACTOSIDASE6 tool Col-0 a MSC than mean the mucilage of necessarily adherent natural not non in does more mutation extrusion even mucilage releases adherent Rak-1 of for absence extrusion the mucilage of al. loss et complete a reach can .thaliana A. MUM2/BGAL6 PRX36 04 Voiniciuc 2014; 07)o h oso deetmclg xrso o a- (Saez-Aguayo Rak-1 for extrusion mucilage adherent of loss the or 2007a) tal. et tal. et tal. et 08.In 2018). noeezmsncsayfrpoe edmclg yrto n xrso (Dean extrusion and hydration mucilage seed proper for necessary enzymes encode utvr (Liu cultivars 03 Saez-Aguayo 2013; tal. et oeo h atrrgltr eogn oteMCtobxas euaetefrainof formation the regulate also toolbox MSC the to belonging regulators master the of some , .thaliana A. 06 u h eeto rsuetwr uiaedspernehglgt t ambivalent its highlights disappearance mucilage a toward pressure selection the but 2016) tal. et (Soto-Cerda 07,adtese abnpriinn (Golz partitioning carbon seed the and 2017), .thaliana A. .thaliana A. o k1 hsrpeet o ubro ee scmae oteMCtobxsize toolbox MSC the to compared as genes of number low a represents This Sk-1. for tal. et 09.A o-yoprosseshv uhbte uonyecec n since and efficiency buyoancy better much a have seeds non-myxospermous As 2019). tal. et tal. et 06 hw rdeto uiaeaudneadmxsem ffiiny This efficiency. myxospermy and abundance mucilage of gradient a shows 2016) tal. et n hti a eue oivsiaewehrfntoa rhlg r present are orthologs functional whether investigate to used be can it that and ETNEHLSEAEINHIBITOR6 PECTINMETHYLESTERASE tal. et 09.Teerslshglgttefc htteMCtobxsat ob well be to starts toolbox MSC the that fact the highlight results These 2019). 09.Uo h addts w ee eeietfid n led known already one identified, were genes two candidates, 8 the Upon 2019). hog eypeieadmlclrpeoyig WSrvae ny8 only revealed GWAS phenotyping, molecular and precise very a through , ouain eae omxsem ovret he ontemgenes downstream three to converge myxospermy to related populations tal. et 06 Miart 2016; 08 loe h dnicto ftesaitclyms eeatsingle relevant most statistically the of identification the allowed 2018) 03 n MI n R3 ucin r ihl n sequentially and tightly are functions PRX36 and PMEI6 and 2013) ( MUM2/BGAL6 tal. et tal. et .thaliana A. 09,o in or 2019), 5 07;Saez-Aguayo 2007a; o h,and Sha, for ) .thaliana A. aua ouain uha h (Macquet Sha as such populations natural tal. et .thaliana A. tal. et 08 Li 2018; tal. et 2014). tal. et ( aua ctps(Saez-Aguayo ecotypes natural PMEI6 tal. et PEROXIDASE36 04 Voiniciuc 2014; tal. et tal. et (Fabrissin PMEI6 04.Ufruaey no Unfortunately, 2014). MUM2/BGAL6 04.Alcharacterized All 2014). o Dja, for ) .usitatissimum, L. 08 hn&Wang & Chen 2018; 2014) , MUM2/BGAL6 tal. et . MUCILAGE- Interestingly, tal. et .thaliana A. tal. et ( 09 and 2019) PRX36 natural 2016). 2007; out L. ) Posted on Authorea 20 Apr 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158739590.04450109 | This a preprint and has not been peer reviewed. Data may be preliminary. and species italica from Brassicaceae two TTG1 the of in demonstrated and Koyama, 2009) been Hiratsu, Hemleben has Matsui, release 2001; and (Penfield production with (MYB5-MYB23-MYB61) as together MYBs Li such 2005; (TT2) additional Ohme-Takagi MYB three cells & Tanaka one least epidermal (TT8-EGL3), at (Golz to bHLHs of in two linked need (reviewed least often (TTG1) at involves WDR process pigments one biological coat the new seed Indeed, and of 4). (Sullivan mucilage events (Figure emergence establishment duplication the mucilage recent control seed and to numerous to co-opted subjected been been have mucilage proteins seed al. of MYB presence and the bHLH of independent The is conserved embryo allowing the is regulator and master SK11/12 coat this by seed for phosphorylated the (Li regulation between Webb be ancestral flow Brockington, can this carbon that Hearn, that in suggesting (Airoldi, 215 switches orthologs, function serine TTG1 ancestral the plant Interestingly, clock essentially seed in 4). circadian across differentiation its Figure cells a to 2019, epidermal undergone Glover addition of has & in control It for plants, plants). functions of (seed new organs gymnosperms acquired all and and that angiosperms node reveals of this family at ancestor factor duplication common transcription (WDR) the repeats this in domain Altogether, appeared WD40 TTG1 2019). the functions. of Nguyen several analysis & control phylogenetic Tran plant actors Land Nguyen, molecular 2015; few Lepiniec a & how Dubos explains epigenetic (Xu, probably or occur post-translational also competitions, may interactions, with modifications through correlates mechanisms that regulation additional species wever, Medicago natural (Li two coat in seed the in GL2 production pigment flavonoid regulator and downstream the of 3) transcription the decreasing plex, (TT2/MYB123) TESTA2/MYB123 TRANSPARENT (Galway with MSCs interaction the in the one type mucilage Accordingly, only wild accumulate 3). controls not Figure mostly H¨ulskamp each MYB 2019; does & (ii) each and traits, Schrader (iii) aforementioned and Chopra, all traits Zhang, ttg1 to more 2019; common liana or Wang (Figure is two TTG1 & of complexes (i) regulation (Chen regulatory knowledge, the trait (MBW) our in of involved to repeat regulation is that, the protein MYB-bHLH-WD40 note bHLH allows to the factor important constitute is transcription They It (WDR) 3). traits. repeats these domain WD40 of a each (TTG1), 1 GLABRA TESTA utgn aiysc sbLsadMB.Teedpiae ee hnfntoal iegda shown as for diverged especially functionally toolbox, then MSC genes the duplicated to These belonging MYBs. genes and other bHLHs for as node such Brassicaceae family multigene the at events (Mabry duplication duplication the whole-genome with a undergone level, have (Doroshkov angiosperm (Doroshkov Brassicaceae the within gene at triplication each hair, resolved for (root well genes, early traits similar is bHLH branching complex-associated a genes TT8 MBW gene Using two all 4). and these ortholog H¨ulskamp Additionally, for (Figure & of 2019). EGL3 (Zhang plants phylogeny angiosperms proteins mucilage) seed and tested the bHLH pigments of all proanthocyanin two and ancestor in anthocyanin the common trichome, demonstrated of recent been conservation most also functional the have the in TTG1 approach, of trans-complementation presence the for al. . tal. et 09 Doroshkov 2019; 07.Ti ucinlcnevto oehrwt la rhlgu eainhp iemr confidence more give relationships orthologous clear with together conservation functional This 2017). h osqec o h ed stepooino ii trg nteeby tteepneo mucilage of expense the at embryo the in storage lipid of promotion the is seeds the for consequence The UIAEMDFE4RANS ISNHSS2 BIOSYNTHESIS MODIFIED4/RHAMNOSE MUCILAGE spoal epnil o h ieeta aac ewe edlpdadpgetmclg contents pigment/mucilage and lipid seed between balance differential the for responsible probably is a etr uiaewl yepeoyeof phenotype type wild mucilage restore can 08 Airoldi 2018; .thaliana A. uatlcsro ar n rcoe,hsardcdlvlo nhcaisadproanthocyanins and anthocyanins of level reduced a has trichomes, and hairs root lacks mutant EGL3 aelasinensis Camellia , tal. et rbsalpina Arabis GL3 h hshrlto fTG ySAG-ieknss1/2(K11)peet its prevents (SK11:12) 11/12 kinases SHAGGY-like by TTG1 of phosphorylation the , tal. et 09.Terdffrn obntoswti eea B opee aeprobably have complexes MBW several within combinations different Their 2019). and 2019). MYC1 tal et (Chopra (Liu 09 iue3.Tefntoa osraino T1i h mucilage the in TTG1 of conservation functional The 3). Figure 2009, . ro arrgltr)oiiae rma neta HHgn fe a after gene bHLH ancestral an from originated regulators) hair (root tal. et tal. et tal. et tal. et 08.Frtepoe salsmn fmxsem,teei a is there myxospermy, of establishment proper the For 2018). 08)o vnfo h oooyeoosspecies monocotyledonous the from even or 2018b) 04,adin and 2014), 09 iue4.Bascca,a ayohrpatclades plant other many as Brassicaceae, 4). Figure 2019, tal. et .thaliana A. .thalianattg1 A. 6 GL2 tal. et 09.Tu,i ilb o upiigt n several find to surprising not be will it Thus, 2019). tal. et tal. et xrsinlvl(Song level expression 08 iue3.Ti ieeta euainof regulation differential This 3). Figure 2018; .truncatula M. 09 iue4.In 4). Figure 2019, 94 Walker 1994; GLABRA2 B euaoycmlxcnrligseed controlling complex regulatory MBW ( MUM4/RHM2) uattruhtercvr of recovery the through mutant , Y ebro h B com- MBW the of member MYB a ( tal. et GL2 (Pang atil incana Matthiola tal. et (Li ) 99 etr 01.In 2001). Western 1999; eeepeso (Liu expression gene .thaliana, A. tal. et moel trichopoda Amborella 07 iue3.Ho- 3). Figure 2017; tal. et 09.Orthologs 2009). 08;Figure 2018); Desl& (Dressel h three the .tha- A. Setaria GL2 et et Posted on Authorea 20 Apr 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158739590.04450109 | This a preprint and has not been peer reviewed. Data may be preliminary. xedtecnett l yoisoosseis one-nutvl,mclg a eangtv regulator (Figure to of negative difficult germination germination a it achene and be the making can for dispersal species, mucilage also in Counter-intuitively, related seed species. germination closely influence myxodiasporous seed between all to different of to completely expected concept be the is can extend myxospermy roles these as However, such 5A). adaptation germination seed and A dispersal seed Yang on biotic and Influence 2012; abiotic 4.1. Western the the in and in mucilage (reviewed pressure seed between germination selection interactions and positive potential for dispersal a constraints. looked under seed recently more is on and community trait influence 2012c), scientific this the its properties, that hydrophilic investigated implies and presence and first adherent its mucilage functions to metabolism, According plant major species. mother displays myxodiasporous the probably for costly it is that establishment mucilage mucilage seed seed that of Considering function ecological in confer insights can New it constraints. why newly of understand the changes kind to present which helpful rapid we to be and part, can and following strong advantage that the the selective myxodiaspory In to a of variability. evolution, functions intra-species ecological during in putative complex existing identified actors MBW direct the downstream of area and of conservation flax dark great MSCs remaining the for A comparing models MSCs. emerging two the the the Altogether, in comparing by mucilage to scale and seed related larger Rosids closely as a the species at specialization such between Brassicaceae deeper zones between located for studies is seed comparative members evolution different mucilage MYB in seed the of in trait concerning recruitment flow each ternary versatile carbon a control balance and through to functions to numerous regulatory appeared with more first allowing modularity, members TTG1 bHLH complex evolution, with plant interacted progressively seed It during tissues. complexes. that MBW suggest the results of These evolution competitive the non-binary characterize of fully depend and complexes of to size MBW helpful hypothesis small in association the the MYB to (Zhang and reinforces due bHLH interaction the this hypothesis, since form, (Doroshkov and this clade, current solve genes comfort its to Rosid MYB in to difficult However, the resolved of 4). variability well of (Figure sufficiently downstream great clade basis not the some Rosid the is for entire phylogeny at the even family in species, is MYB myxospermy two ancestor of the origin common between ancestral last to shared functions in and their orthologous implicated sequences As all genes in genes. seven conservation flax, of a (GWAS) in show study content (Figure 2018), association species mucilage MBW Rosids wide the the seed genome that among by suggesting conserved the are 2014), identification pigments Dixon the coat & Furthermore, seed Jun and 4). (Liu, mucilage mucilage seed to and 1971). dedicated pigment complexes coat Whitehouse seed & the regulate (Vaughan more positively Brassicaceae they the in as and genes, such complex, MYB MSCs, Concerning MBW of ( myxospermy-related diversity network morphological the the observed of in the part expected downstream of the be in can are genes changes intra-Brassicaceae few an family, indicates this Rosid mutant, triple of same morphology the of both in transcomplementation phenotype between mucilage the divergence the by complement mucilage to seed able gl3 of thaliana rescue partial A. unexpected the by .alpina A. .thaliana A. / egl3 tal. et / etcru scopulophilus Leptocereus tt8 MYB5 Ssa oprdt hs of those to compared as MSCs 09 n fps rnltoa oictos(Li modifications translational post of and 2019) GL3 S olo ee emt aeudroea smerclslcinpressure selection asymmetrical an undergone have to seem genes toolbox MSC rpemtn Zag&Husap21) ic h rbdpi L snot is GL3 Arabidopsis the Since H¨ulskamp 2019). & (Zhang mutant triple ee rmterls omnacso,i gemn ihtecontrasted the with agreement in ancestor, common last their from genes and reii monosperma Artemisia TT2 rhlggnsin genes ortholog tal. et Bris lrs ozae-ors&Plaoa21)and 2015) Palmarola Gonz´alez-Torres & Flores, (Barrios, e.g. 7 09.Snetepyoeyo hsmlieefml is family multigene this of phylogeny the Since 2019). HH) h ihrtedvrec ilb responsible be will divergence the higher the bHLHs), .thaliana A. .thaliana A. Hag&Gtemn19a.In 1999a). Gutterman & (Huang .truncatula M. .thaliana A. S olo ee (Soto-Cerda genes toolbox MSC (Chopra .thaliana A. aeacnevdfnto because function conserved a have tal. et .thaliana A. tal. et uhas such 08,mr tde ilbe will studies more 2018), hscudb ovdby solved be could This . 04.Smlry neach in Similarly, 2014). .apn L nthe in GL3 alpina A. aeiasativa Camelina . lpai persica Blepharis tal. et tal. et or , Posted on Authorea 20 Apr 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158739590.04450109 | This a preprint and has not been peer reviewed. Data may be preliminary. olpyia rpriscnhv ao mato ae viaiiyadro eerto Fgr 5B- (Figure penetration root and availability water on period. impact drying major long a a following have flooding natural can or more properties cycles, in drying role physical its and Soil investigating wetting necessitate of compared would succession as management as water ways, such in dissemination (Arshad mucilage situations buoyancy seeds seed and imbibed of run-off fully roles water and complex seeds wind, imbibed (Lenser for never dehydration seeds, to re-dried upon Kovalev the size (Kreitschitz, for their properties compromise, and dispersal structure its least mucilage their In even at seed conserve 2016). and influencing the 2015, time level of hydration longer Gorb properties its for & frictional to surface and according adhesive water Interestingly, change on 1999b). can float Gutterman can & that mutants (Huang of seeds removed or Achenes type myxospermous it. wild non on Saez-Aguayo the to than germinate 5B-3; compared faster as (Figure seed water water their residues imbibed provide in (Saez-Aguayo acid mucilage, may mucilage of galacturonic lack it un-released a with that with have linkages that propose mutants ionic to Indeed, through 2014). tempting it is sequesters it but gels, For embryo. hydrophilic the highly the for constitutes by mucilage seeds seed attached As constraints two abiotic their facing dehiscence, and its Influence fruit after 4.2. even the it, of to 2018). sticked halves seeds Erbar two four & the the Fligge keep between (Leins, to for fruit wind dispersion place the optimal differential of dispersion. an surface a until its inner allowing dispersal the stops efficient from Interestingly, and it an 5A-2). secreting by water (Figure by advantage encounters reached powerful is a seed favors give conditions the anemochory can hydric traits while until plant both conditions of mother dry Combination the under from of dissemination far seeds dispersion wind-driven on seed prevents wings Antitelochory and complementary. mucilage seed deserti of phyton characterized co-occurrence having The delay. species Brassicaceae Toorop three 2010; For 2018a). namely persistence Song myxospermy, arabicum, species improve & for to Tanveer seeds (myxosper- seeds Wang, species dimorphic dimorphic same (Liu, these the use within dispersion species occur and morphotypes These seed seeds). two non-myxospermous water that and to fact Therefore, the mous permeability of 2016). seed advantage and/or Gallacher took uptake studies & water Some Santo to related (Bhatt, gazes. be effects to to possibly germination seems and germination contrasted in to function led mucilage the removal its amplexicaulis), but Plantago and uptake ciliata Plantago elegans, Boerhavia eedn etrgriainpeoye r bandwt E plcto n o ihsl tequal at salt with not and application PEG with (Geneve obtained (Geneve mucilage are potential phenotypes without osmotic germination achenes better the dependent than better germinate in germination the on in mucilage (Arsovski of obtained rate effect were germination positive on a than suggesting germination, rather for delayed efficiency observed a induce rate extrusion germination mucilage of seed decrease (PEG)-dependent glycol most myb61 polyethylene pioneer the The in 5A-1). considering function (Figure conditions, germination mucilage improve (Witztum, seed also germination Gorai prevent can in cited then mucilage time However, (last and re-emphasized demonstrated. excess regularly fully water was never under function This transfer 1969). oxygen Evenari block & Gutterman could mucilage seed the , .thaliana A. , gl2 h edmrhtp ihtehge omnyaentmxsemu L,Tn akn&Baskin & Baskin Tan, (Lu, myxospermous not are dormancy higher the with morphotype seed the and tal. et emnto eevst edel xlrd For explored. deeply be to deserves germination (Gorai ttg1 02 Arshad 2012; .thaliana A. uat Pned20) naohrMCtobxdwsra eemtn,adfc in defect a mutant, gene downstream toolbox MSC another On 2001). (Penfield mutants mum2 tal. et .arabicum A. tal. et 04 usin hte nieohr n nmcoyaeopst rcnbe can or opposite are anemochory and antitelochory whether questions 2014) reii sphaerocephala Artemisia or myb61 .thaliana A. tal. et h edmclg ae ag muto ae rmteenvironment the from water of amount large a takes mucilage seed the , 07.Uo v eetseis( species desert five Upon 2017). h edmclg hc br mriguo miiinaeal to able are imbibition upon emerging fibers thick mucilage seed the , 09 ugsigta yoprossessol emnt without germinate should seeds myxospermous that suggesting 2019) uat (Saez-Aguayo mutants tal. et itohru tits aslabrapsoiiadAethionema and bursa-pastorisi Capsella strictus, Diptyocharpus a oiierl uiggriainudromtcstress osmotic under germination during role positive a was 04.Hwvr h edmclg losfs edsinking seed fast allows mucilage seed the However, 2014). tal. et 8 emnt n otbte hntermclg was mucilage their when better float and germinate tal. et uai annua Lunaria 09.Hwvr oercnl osc phenotypes such no recently more However, 2009). .hispanica S. 09.Amr opeeudrtnigo the of understanding complete more A 2019). tal. et tal. et h edmclg rsneicesdwater increased presence mucilage seed the tal. et .strictus D. aadl und,Lpdu aucheri, Lepidium subnuda, Lavandula 04.Tu,terl fse mucilage seed of role the Thus, 2014). 07.Itrsigy edmucilage- seed Interestingly, 2017). 06 loigadseslefficiency dispersal a allowing 2016) hw upiigueo mucilage of use surprising a shows h natmxcrosachenes myxocarpous intact the , (Lu tal. et tal. et 00 and 2010) 04 though 2014) Heno- tal. et Posted on Authorea 20 Apr 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158739590.04450109 | This a preprint and has not been peer reviewed. Data may be preliminary. oia iest ae tdffiutt rc akteeouinr rgno edmclg ae nyon only based mucilage seed of origin evolutionary morpho- the features microscopic back microscopic striking trace characterized of This diversity deeply levels. to available surprising inter-species difficult The a and it morphology. from intra-species makes results the diversity that both logical trait at seed angiosperms, macroscopic among a is Myxodiaspory perspectives functions and characterized previously Conclusions the the affect to could compared that as functions manners astonishing to unexpected revealing germination uncovered in from be development development to (Roberts because plant starts plant level protocarnivory interactions nutrient increases of biotic low mucilage case in under seed involvement a especially in be establishment, nematodes to seedling plant trapped seems young of a it Surprisingly, death than 2018). massive rather Provan the adaptation & Warren plants. nematode for (Roberts, the organisms nematodes dangerous to more due even of probably predators bursa-pastoris, more be is could nematodes attraction Conversely, adaptation. this (Tsai nematodes, proteins development. those and plant of on carbohydrates impact seed-surface pathogenic of major have requirement can and plants thaliana with A. (Raviv interaction close independent even in are are species mucilage Nematodes several seed and of secretion coat protein seed that the suggesting from species secreted (Raviv myxospermous are old mucilage, non chitinase, decades with several and mycor- effect seeds protease, arbuscular positive in (Hu nuclease, for combined establishment as significant responsible seedling such show fungi enhance enzymes, not to these independently do of probably plants) growth acting with seedling symbiosis on patho- (mutualistic effect and same rhizae positive drought favor the the from to in However, seedlings diversity young mucilage. and explained protecting composition recently community activities, was (Hu microbial enzyme effect gens soil soil beneficial on and effect This interaction positive fungal-bacterial 2012b). mucilage Huang the through & CO species Zhang providing Baskin, micro-organisms, Baskin, by (Yang, microbial degraded promoting be 5C-5) to (Figure effect” media with “selective nicola illustrated This further 2010). was development Schrempf hyphae & to (Meschke seeds, comparison on in symptoms mucilage co-inoculated se are seed micro-organisms the within both When proliferation in constraints wilt. shown biotic verticillium was with the plant causes dealing the for in organs, community mucilage microbial plant of lividans seed on Streptomyces around importance of influence involvement omnipresence the Its an their 5C). consideration indicate and (Figure into could development Taking investment their environment. metabolic along this the physiology in plant released in (Knee micro-organisms polysaccharides mucilage of cell amount border cant root to Similar (Figure soil of interactions kinds Biotic diverse 4.3. as in such days species 30 of least lifestyle at non-disseminating for the stability with aggregate increase Marsico Di to 5B-4; from particles extract soil mucilage links seed which (Deng adding time By longer 2012a). for Huang water & pastoris Liu bursa Baskin, C. Baskin, (Yang, mortality plant reduces of myxocarpy The 4). ug (Geneve fungi tal. et edmclg otiue oatatro-ntnmtds(iue5-)wt h additional the with 5C-6) (Figure nematodes root-knot attract to contributes mucilage seed lsl eaieseisof species relative closely a 09) lmrmct soeo h ruso uglseispstvl matdb achene by impacted positively species fungal of groups the of one is Glomeromycota 2019b). tal. et olrelgclpoete r oie atclryfrhdalccnutvt retaining conductivity hydraulic for particularly modified are properties rheological soil , tal. et htihbt emnto n rwho h ahgncfungus pathogenic the of growth and germination inhibits that 08.Teeoe edmclg ol oal mrv h olrelgcli agreement in rheological soil the improve locally could mucilage seed Therefore, 2018). reii sphaerocephala Artemisia ahnssativus Raphanus 07.Tedsr plant desert The 2017). tal. et 04.Asmlreeti rvkdby provoked is effect similar A 2014). tal. et 07.Hwvr hsaiiyt ert rtisi osre nthe in conserved is proteins secrete to ability this However, 2017). .thaliana, A. tal. et ri the in or avahispanica Salvia nacsseln mrec nissnyevrnetand environment sandy its in emergence seedling enhances .thaliana A. 01,nnahrn edmclg osiue signifi- a constitutes mucilage seed non-adherent 2001), 2 reii sphaerocephala Artemisia 9 n oul uast rmt edigestablishment seedling promote to sugars soluble and .thaliana A. a yoprossesta r loal oattract to able also are that seeds myxospermous has .dahlia V. .tain gl2 thaliana A. tal. et uiaesceoycl MC olo genes toolbox (MSC) cell secretory mucilage ceemclg and mucilage achene n osdrbyrdcdtepatdisea- plant the reduced considerably and xrtn o-deetmucilage. non-adherent excreting 09.Cnieigteprstcnature parasitic the Considering 2019). tal. et .hispanica S. uatdpie fse mucilage seed of deprived mutant tal. et 09) ciestress-associated Active 2019a). tal. et .thaliana A. ceemclg a shown was mucilage achene 08.Tu,se mucilage seed Thus, 2018). 2017). etclimdahlia Verticillium edmclg addition, mucilage seed .lividans S. olttihmgrami- Colletotrichum sn h bacteria the using a better a has Capsella that Posted on Authorea 20 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158739590.04450109 — This a preprint and has not been peer reviewed. Data may be preliminary. h relae r h nsuei h eiwdsuis G3 steacso eeo tG3 tL and AtGL3 AtEGL3, of gene ancestor the is EGL3* to studies. specific reviewed be the in to use appear ones box the tool are leaves MSC tree the the of actors enzymatic evo- downstream sequential in the trait of overview myxospermy studies an characterization giving functional plants and land the of of tree lution phylogenetic Simplified spatiotemporal (Golz 4: the release Figure of and regulation production the 2019). mucilage together Wang enables (P), including interaction & serine traits Chen competitive conserved seed of a various modularity of of MYB status specificity phosphorylation and the bHLH and a (TTG1) with par- member carbon WDR spatiotemporal regulator the master regulate to complexes in cell (MBW) from titioning MYB-bHLH-WDR expending The domains wall 3: primary Figure of and rupture (am) pollination mucilage sequential after adherent and DAP=day between m4) rup- distinction cell. to simultaneous the (m1 the and in layers columella, cells features mucilage in MSC volcano-shaped different Major features each a (nam). Major in of layers domain mucilage presence models: wall non-adherent the both cell with between primary dynamics differences of wall numerous ture cell the complex Note highly scale. : to drawn not are al. (MSCs) model cells historical secretory the mucilage Francoz seed in of development deve- localization MSC (MSC) the (A) mucilageof cell illustrating imbibition. polysaccharidic magnification secretory water and intracellular mucilage view upon myxospermous dynamics, wide seed mechanisms in wall for extrusion found cell and species commonly of organization, model diversity is as the fruit flax illustrating per lopment and seed Arabidopsis one 2: than Figure more that secretory and mucilage scale species. of to myxocarpous species. drawn origins vs not epidermal myxospermous are two in layers release the mucilage comparing enabling scheme cells Developmental 1: language. Figure english the (ANR-18-CE20- improving “MicroWall” for Uhlmann LEGENDS: project Charles FIGURE (ANR-10-LABX-41; thank Agency to “TULIP” Research III. like project Sabatier-Toulouse National would Excellence Paul We French of University 0007). Laboratory the the French by the and their by funded supporting ANR-11-IDEX-0002-02) supported scholarship for also PhD CNRS was a work and from (France) This benefited III S.V. Sabatier-Toulouse Universit´e work. Paul to research thankful are new authors uncover The to tracks abiotic promising range and most wide the biotic environments. probably the particular The ACKNOWLEDGMENTS and to to environments. field related contributes contrasted this functions face probably in integrated that points traits mucilage studied seed of species least diversity each the this in or are pat- patterns turn, observed constraints various sub-layering In functions origins, and ecological release. epidermal composition mucilage of polysaccharidic MSC mucilage two of times various including several modes dynamics, evolved traits wall several have diverse cell may MSC highly and mucilage of of evolution complex, as terns combination plants regulatory toolbox seed a this MSC From during as the Rosids. sequentially independently across of appeared mucilage conserved genes complex that shed notably downstream regulatory pressure is to master few selection starts upstream a intense which MBW on the myxospermous related species, applied hand, non between mainly one in or be the illustrated to species On appears a story. undergoes within evolutionary establishment actors mysterious this molecular on these light of comparison allow now 09.Te30-50 The 2019). tal. et 09,adi h mrigmodel emerging the in and 2019), rbdpi thaliana Arabidopsis .thaliana A. μ .thaliana A. ieclsfo ohseisaedant iia cl,terlae uiaelayers mucilage released the scale, similar to drawn are species both from cells wide m S olo sn ulse hlgnmc transcomplementation phylogenomic, published using toolbox MSC eunilyeovdadwr opeea h oi oe n htthe that and node, Rosid the at complete were and evolved sequentially .thaliana A. ed hog ope modularity. complex a through seeds rbdpi thaliana Arabidopsis oeta h ebr fteMWcmlxcnrligthe controlling complex MBW the of members the that Note . aua ouain.O h te ad h edmucilage- seed the hand, other the On populations. natural iu usitatissimum Linum 10 .usitatissimum L. ntelf aatdfo Wsen2001; (Western from (adapted left the on ntergt(dpe rm(Miart from (adapted right the on eunilsnhsso orhighly four of synthesis sequential : .thaliana A. ocpulse rs section cross seed conceptual tal. et h osre ancestral conserved The oeta h different the that Note 08 Li 2018; pce ae on names Species . (B) tal. et .thaliana A. Kinetics 2018; et Posted on Authorea 20 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158739590.04450109 — This a preprint and has not been peer reviewed. Data may be preliminary. ossMM eeecdsa-aatsds eurdfrtepouto fse otmclg ihcorrect Arabi- with The mucilage (2007) coat G.W. seed Haughn of production . . the . for Y.T., required Hwang -Galactosidase D.S., properties. Young a hydration J., encodes Huang gene H J., MUM2 and Tewari dopsis . H., . determination . Zheng M.C., fate G.H., Albani Dean cell G., Coupland epidermal in S., for Schellmann function J., TTG1 regulator Span of five H., common Wolff in a D., germination Chopra GLABRA2, and (2019) uptake Arabidopsis. water S. in on biosynthesis Wang anthocyanin effect & mucilage S. Seed Chen (2016) regulating desert. pathways Arabian D. hyper-arid interacting Gallacher the of & from network A. species complex Santo the A., of Bhatt organization. part matrix as polysaccharide function, mucilage 2 seed G COBRA-LIKE Arabidopsis I., of Ben-Shlush germination role A., the The Hugger in A., mucilage Idan-Molakandov of D., role Ben-Tov The (2015) A. Palmarola & Gonz´alez-Torresof L.R. J., in Flores success D., reproductive Barrios underlying innovation evolutionary Seeds—An (2019) plants. U. flowering Grossniklaus in AtBXL1 & modification (2009) C. arabinan T.L. Baroux Western pectic & for cells. M.C. required secretory McCann mucilage -L-Arabinofuranosidase N.C., Arabidopsis -D-Xylosidase/ Carpita bifunctional G.W., a research. Haughn wall encodes T.M., cell plant Popma for A.A., model Arsovski a As (2010) T.L. annual Western Behavior & the & G.W. in Haughn Mummenhoff diaspores a., & A. dimorphic G. 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R. 261-276. , ln Science Plant rnsi ln Science Plant in Trends ln n Soil and Plant aslabursa-pastoris Capsella 232 109 avahispanica Salvia 1–11. , 419–427. , ln n Soil and Plant 272 rbdpi thaliana Arabidopsis 179–192. , 374 727–738. , 42 aslabursa-pastoris Capsella . mrvssi grgt stability. aggregate soil improves L.) eohtndeserti Henophyton 387 591–605. , 20 rpScience Crop .Mdk sehr’ us)o olwtrrtnin tblt and stability retention, soil-water on purse) (shepherd’s Medik. L. 167–176. , 515–524. , .tihm development. trichome L. 12 57 e Phytologist New 2160–2169. , neatv ffcso eprtr,slnt and salinity temperature, of effects interactive : sehrspurse). (shepherds 225 e Phytologist New ln n Soil and Plant M ln Biology Plant BMC ln Biology Plant 1461–1469. , eeomna Biology Developmental 11 214 425 ln Physiology Plant 204–212. , 959–966. , 57–69. , Developmental 19 ora of Journal Matthiola 53. , Posted on Authorea 20 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158739590.04450109 — This a preprint and has not been peer reviewed. Data may be preliminary. i . hmYY,Se . agY,GohS enyMJT 21)Vraino opsto and composition of Variation (2016) M.J.T. Reaney & ( S. flaxseed Technology Canadian and Ghosh Science six Y., Food from of Wang gum Journal J., of Shen properties biosynthesis Y.Y., polymer functional coat Shim seed in J., roles pivotal Liu play MYB14 Arabidopsis and MYB5 The (2014) (2009) R.A. R.W. Dixon in & Parish J.H. Jun . . C., . Liu J., morphogenesis. Preston trichome R., and Cell development, Napoli coat Plant seed TESTA R., The synthesis, TRANSPARENT mucilage Seyit of regulates H., factor phosphorylation transcription Pham MYB5 Site-specific O.N., (2018) Milliken H. S.F., seeds. Yu Li Arabidopsis & in L. partitioning Ji carbon B., mediates Chen GLABRA1 B., Zhang C., Li in dimorphism seed and 172 fruit of (Brassicaceae). plasticity and Lunaria control Cevik of Developmental K., anemochory Graeber in T., sails Lenser as valves Silique (2018) C. Erbar Biology & K. Fligge P., Arabidopsis. Leins in extrusion Spatiotemporal mucilage (2013) coat I. Hara-Nishimura seed 1355–1367. & for , K. required Nishitani is M., PEROXIDASE36 Nishimura of M., secretion Kondo T., mucilage. functi- Shimada coat of 2 T., seed SAUCER Kunieda properties FLYING normal Arabidopsis establish physical (2019) to G.W. FLY1 the Haughn with & – redundantly T. ons invasion” Demura I., “Sticky Hara-Nishimura T., (2016) Kunieda S.N. 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Northwest of of Germination areas (1999b) desert Y. sandy Gutterman of & concentrations. achenes composition Z. salt mucilaginous and Huang by by coat affected absorption as Water seed germination (1999a) the and Y. of floating Gutterman Differentiation & (2015) Z. Huang H. Lan & J. Sinica Cao Biophysica of J., mucilage Yuan the C., of Wang D., Huang eiaotruncatula Medicago 1691–1707. , 9–22. , 20 .se mucilage. seed L. 238–243. , 17 21 iu usitatissimum Linum 152–159. , 47 eiimperfoliatum Lepidium 72–89. , 14 775–787. , . ln Physiology Plant 775–784. , elti ora fNanotechnology of Journal Beilstein .. Adig O.S., ¨ .se uiaiosevlp n t ilgclsignificance. biological its and envelope mucilaginous seed L. ot fia ora fBotany of Journal African South zlN,D N., uzel ¨ 51 . eetana ihtpclmyxospermy. typical with annual desert a L.: 165 2313–2326. , LSONE PLOS 1424–1439. , lr:Mrhlg,Dsrbto,Fntoa clg fPlants of Ecology Functional Distribution, Morphology, Flora: ne .. rsh . ebe-eze .(2016) G. Leubner-Metzger . . . C., Grosche A.A., ¨ onmez reii sphaerocephala Artemisia 13 13 iu usitatissimum Linum salJunlo ln Sciences Plant of Journal Israel e0200522. , aueCommunications Nature 7 1918–1927. , ln n elPhysiology Cell and Plant ehoeaarabicum Aethionema 65 naso botany of Annals 187–196. , Atrca) curn nthe in occurring (Asteraceae), . cultivars. L.) reii monosperma Artemisia 9 caBohmc et Biochimica Acta h ln Cell Plant The 571. , . oeua Plant- Molecular ln Physiology Plant 110 47 61 International 27–34. , 205–212. , 308–317. , Plantago Plant Acta 25 : Posted on Authorea 20 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158739590.04450109 — This a preprint and has not been peer reviewed. Data may be preliminary. efil .(01 Y6 srqie o uiaedpsto n xrso nteAaiossse coat. seed Arabidopsis WD40 the A in (2009) extrusion R.A. and deposition Dixon mucilage for . Cell required . . Plant is The MYB61 D., (2001) Huhman S. J., Penfield development. Wen trichome for G.J., not Peel but biosynthesis K., from Saathoff variants. protein J.P., natural repeat Wenger of exploitation Y., the Pang and for production perspectives polysaccharide future Understanding (2014) mutants: M.C. Botany coat Ralet seed & using S. Saez-Aguayo properties A., epigenetically Berger (MBWi) of H.M., interferers North structure MYB–bHLH–WDR The molecular targets. (2019) and MBW’s N.H. composition Nguyen the chemical & suppress G.B. on Tran study Cytological C.T., gums. Nguyen review (2019) seed A K. and exudates Pageau (2012) gum B.T. plant Amid . newly . . & mucilage. L., H. flax Dupont Mirhosseini of H., nature layered Demailly the E., reveals Petit analysis Science chemical N., with Dubrulle combined F., approaches Fournet seed F., on Miart mucilage of deposition the (2010) of dahliae H. repressor Schrempf inhibition AtMYB23 & and chimeric H. Meschke stems, A (2005) and M. leaves Ohme-Takagi Arabidopsis. of & in elongation and H. coats Tanaka chemical roots, T., hairy situ, Koyama induces In K., Hiratsu (2007b) K., H.M. Matsui North & A. of Marion-Poll composition ology J., the of Kronenberger study (2007a) M.C., macromolecular H.M. Ralet North a mucilage. A., & affects seed Macquet A. in accession Marion-Poll I Arabidopsis G., rhamnogalacturonan an Phylogeny of Mouille in potential (2019) J., mutation hydrophilic C. Kronenberger occurring Pires O., naturally Loudet . . A M.-C., . Brassicales. C., Ralet the Bottoms A., in W., Macquet events Dismukes duplication B., whole-genome Sutherland independent P., annual multiple Blischak desert and J., cold Brose the M., in Mabry heteromorphism seed and Fruit (2010) 999–1014. C.C. , Baskin & in J.M. Advances Baskin co-option ephemeral (2017) and D., A. Tan factor from J., Gonzalez WRKY regulation. Lu protein & a betalain A. of repeat for Cantero addition MYB WD40 A., model: anthocyanin A regulatory Bean an pigment (2018b) A., of (MBW) Campbell T. Repeat L., Xia MYB-bHLH-WD Aguirre the . A., . . Brockman A., W., Lloyd Chen MYB– of X., formation Dai the complexes. through P., ternary accumulation Wang proanthocyanidin bHLH–WD40 and X., anthocyanin regulates Jiang sinensis H., Camellia Hou halophytes GLABRA of Y., TESTA adaptation TRANSPARENT Liu important an (2017) heteromorphism: Seed M. (2018a) heterogeneity. ( Chen J. habitat Song millet & for . . M. foxtail Tanveer . L., to Wang S., R., Arabidopsis Duan Liu from C., development Gao and C., growth Jin plant ). D., regulates Li ubiquitously S., 1 Qi K., Liu ln Science Plant 48 nsesadrosof roots and seeds on 114 10 984–999. , itcoapsstrictus Diptychocarpus 684. , 1251–1263. , 13 254 2777–2791. , eiaotruncatula Medicago 60–69. , ln n elPhysiology Cell and Plant rnir nPatScience Plant in Frontiers ilg fteCell the of Biology rbdpi thaliana Arabidopsis nentoa ora fMlclrSciences Molecular of Journal International tetmcslividans Streptomyces Bascca)adpsil dpiesignificance. adaptive possible and (Brassicaceae) sncsayfrtsu-pcfi nhcai n proanthocyanidin and anthocyanin tissue-specific for necessary is odRsac International Research Food rbdpi thaliana Arabidopsis ln Physiology Plant 111 ln n elPhysiology Cell and Plant 46 . 147–155. , 9 14 irba Biotechnology Microbial 284–291. , 1515. , niistepoieaino h fungus the of proliferation the inhibits h ln Cell Plant The 151 edca mucilage. coat seed 1114–1129. , β- 46 -aatsds hticessthe increases that d-Galactosidase 58 387–398. , 3 iRi preprint bioRxiv 19 428–443. , 1431–1441. , 19 1686. , 3990–4006. , naso Botany of Annals ln n elPhysi- Cell and Plant rnir nPlant in Frontiers eai italica Setaria Verticillium . nasof Annals 105 Posted on Authorea 20 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158739590.04450109 — This a preprint and has not been peer reviewed. Data may be preliminary. fse omnyadflwrn iei h rbeweed arable the in Co-adaptation (2012) time P.P.M. Iannetta flowering & and G.R. Squire (2018) dormancy B., T. seed Locardi Ishimizu G.S., of Begg R., Cuerva . . Campos . P.E., Toorop K., family. Yagyu glycosyltransferase H., plant-specific Kajiura novel a K., 4 represent Tsuruhama rhamnosyltransferases M., RG-I Ogawa-Ohnishi Pectin K., Kato Y., Takenaka Science Plant industry food in biopolymer emerging as mucilage seed Plant (2018) L. Hoffmann applications. & associ- C. Genome-wide Gaiani (2018) C., ( F.M. Soukoulis You flax & in M. content hull Olivos carbon Sciences and H.A., of Molecular mucilage Gajardo Regulation of R., comparative (2017) analysis Quian a R.J. ation S., orbicularis: Rose Cloutier Medicago & B.J., and M.L. Soto-Cerda truncatula Garg Medicago legume S., model Wheeler the N., of approach. 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S., epidermal D., Saez-Aguayo coat Ropartz seed in L., methylester- homogalacturonan limiting Botran of by release A., ification mucilage Arabidopsis Berger promotes INHIBITOR6 METHYLESTERASE M.-C., general. PECTIN in Ralet myxodiaspory S., on notes Saez-Aguayo with Plants (Lamiaceae) of Nepetoideae Geography the and in in tematics Myxocarpy protocarnivory (2001) facultative O. for Ryding Evidence (2018) J. Provan & pastoris J.M. Warren H.R., of Roberts innovations phylogeny. reproductive unified and a Vegetative dead (2000) for Sciences The D.J. implications Biological (2017) Garbary plants: G. & land Grafi D.L. enzymes. Nickrent early & hydrolytic R.J., active Y. Duff for Gutterman K.S., storage O., Renzaglia long-term Frenkel of a V., as Makover properties functions G., physicochemical coat In Granot and seed L., Composition Aghajanyan mucilage. (2019) accessions. B., seed natural M.-C. Raviv Arabidopsis of Ralet of & structure seeds H.M. and from North mucilage composition outer L., the Botran into D., insights Poulain New (2018) Reviews R.A. Plant Burton Annual & J.L. Phan ulvnAM,Asvk .. hmsnA,SnsrmR,TumnRE,Np . uishC. Queitsch . . . S., Neph maturing R.E., in Thurman DNA R., regulatory Sandstrom of dynamics A., and Thompson Mapping A.A., (2019) Arsovski A.M., Sullivan ˇ ˇ 669–676. , p 0511.Wiley. 1085–1116. pp. . seeds. r nir nPatScience Plant in Frontiers urn pno nFo Science Food in Opinion Current 10 cetfi Reports Scientific 1434. , 355 19 p –1 onWly&Sn,Ld hcetr UK. Chichester, Ltd, Sons, & Wiley John 1–41. pp. . 2870. , 769–793. , ln Cell Plant 71 8 10120. , 8 31 503–514. , 2070. , 809–831. , 22 28–42. , iu usitatissimum Linum 15 LSGenetics PLoS hlspia rnatoso h oa oit B: Society Royal the of Transactions Philosophical aslabursa-pastoris Capsella h ln Cell Plant The o PLANTS AoB rbdpi thaliana Arabidopsis 10 LSONE PLOS . seeds. l.) 25 e1004221. , 11 308–323. , ln elphysiology cell & Plant plz031. , sehrspurse). (shepherds nentoa ora of Journal International nulPatReviews Plant Annual siliques. 12 e0181102. , aslabursa- Capsella auePlants Nature rnir in Frontiers Annals Sys- 59 , Posted on Authorea 20 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158739590.04450109 — This a preprint and has not been peer reviewed. Data may be preliminary. agX,Bsi .. aknJM,LuG un .(02)Se uiaeipoe edigemergence seedling improves mucilage Seed (2012a) cellulose Z. for Huang shrub. & essential desert G. is sand Liu TRM4 a J.M., interacting of Baskin (2019) and C.C., B. organization Baskin Usadel X., microtubule Yang & cortical maintaining H. by Klose mucilage CESA3. S., seed with Dieluweit Arabidopsis L., in Fu deposition C., Voiniciuc during B., barrier MYB-bHLH-WDR Yang by oxygen biosynthesis flavonoid an of control Transcriptional as (2015) L. mucilage complexes. Lepiniec & Integumentary C. Dubos (1969) W., Xu M. Evenari & (2020) of Y. H.M. germination North Gutterman . A., AFM. . . by Witztum Y., visualised seeds An Arabidopsis J., for of Malmstrom Role mucilage V.V., (2018) outer 21 Symonds the T. A.H., in Munnik structures Irani Polysaccharide . V., . . Cornuault A., serration. M.A.K., leaf Guardia Williams and F.R., attachment, Marquez mucilage M., seed Lamers movement, Science X., stomatal in Zarza PLC7 Q., Arabidopsis Zhang germination R., seed in Wijk role van and genetics, production, mucilages: coat seed dispersal. of cell and tale secretory sticky mucilage The (2012) coat T.L. seed Western in defective mutants of Arabidopsis. characterization in and development Isolation (2001) regulates T.L. Western 2 Protein Tubby-like (2019) Botany Y. Experimental Kong of & G. M Zhou K., mucilage. Weitbrecht R., coat Hu seed Y., Arabidopsis Wang in R.I., biosynthesis Ahmed homogalacturonan Gray and Z., protein. differentiation Xu . repeat . trichome . M., WD40 Wang regulates T.L., a which Blundell encodes locus, arabidopsis, N., in GLABRA1 Srinivasan biosynthesis TESTA C.M., anthocyanin TRANSPARENT James The A.C., (1999) Bolognesi-Winfield J.C. P.A., Davison A.R., structure. Walker mucilage seed Arabidopsis in G variation M.H.-W., natural Schmidt E., Zimmermann C., Voiniciuc walls. cell secondary specialized produce 16 cells epidermal G M.H.W., coat mucilage. Schmidt seed B., Yang in C., synthesis Voiniciuc pectin for Cruciferae. enzymes the key of of G tification taxonomy K.A., the Engle and C., Voiniciuc structure Society Seed Dissecting Linnean (1971) (2017) the R.A. J.M. of Burton Journal Whitehouse & . . J.G. . Vaughan M.G., in Hahn biosynthesis heteroxylan N.J., spectroscopy. mucilage Shirley infrared coat K., seed and root- Neumann for of basis J., Regulation genetic Phan (2019) the C., S. Sawa Ma & M.R., attractants. Tucker B. mucilage-derived Favery seed-coat L., by Perfus-Barbeoch behavior C.N., nematode Nguyen knot T., Higaki A.Y.L., Tsai Botany of 1450–1459. , 3452–3473. , 9 1721. , 109 rnsi ln Science Plant in Trends e Phytologist New edSineResearch Science Seed lpai persica Blepharis 481–489. , le .&LunrMtgrG 21)Frto h ak al edgermination. seed early mark: the off First (2011) G. Leubner-Metzger & K. uller ¨ lSone PloS 62 rnir nPatScience Plant in Frontiers 3289–3309. , n . ilwi . cmd ..W,Yn .Y,...Uae .(08 Iden- (2018) B. Usadel . . . J.-Y., Yang M.H.-W., Schmidt S., Dieluweit M., unl ¨ ln Physiology Plant 221 64 . 7 oaia Gazette Botanical 383–409. , e34897. , 881–895. , 20 22 176–185. , 1–25. , n .&Uae .(05 trigt e:HwAaiossseed Arabidopsis How gel: to Starting (2015) B. Usadel & M. unl ¨ 127 998–1011. , n . uL,NrhHM sdlB 21)Extensive (2016) B. Usadel & H.M. North L., Fu M., unl ¨ 130 16 8 328. , 238–241. , rnir nPatScience Plant in Frontiers nentoa ora fMlclrSciences Molecular of Journal International ln Physiology Plant ln oeua Biology Molecular Plant lnaoovata Plantago oeua Plant Molecular h ln Cell Plant The 178 sn am irradiation gamma using 7 12 1045–1064. , 803. , Biomacromolecules 99–112. , 11 rnir nPlant in Frontiers 99 1337–1349. , 421–436. , Botanical Journal Posted on Authorea 20 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158739590.04450109 — This a preprint and has not been peer reviewed. Data may be preliminary. hoX,Qa .&W .M 21)Eetv xrcino rbdpi deetse uiaeb ultrasonic by mucilage seed adherent Arabidopsis of extraction Effective (2017) A.-M. Wu treatment. & L. Qiao X., Zhao thaliana plants. dopsis H in & proteins B. Zhang WDR and bHLH, MYB, 3197–3209. H of & formation A. Schrader complex D., of Chopra mucilage B., Multi-layer Zhang (2017) J.R. Stokes & V. Bulone J., Polymers Stenson structure X., patterned Xing ovata The W., (2018) Plantago mucilage. P. Zeng seed Dupree G.E., in Yakubov . cellulose L., . Yu . to A., bind Li may X., it importance, Yu suggests T., ecological galactoglucomannan Kotake coating: of C.S., Pereira a J.J., Lyczakowski just L., than mucilage. Yu coat More seed (2012c) Systematics of Z. and relationships Huang soil Evolution phylogenetic & by and C.C. mucilage occurrence Baskin seed taxonomic plant. J.M., of Baskin dune Degradation X., sand (2012b) Yang desert Z. a Huang of & growth W. seedling 872–883. Zhang early J.M., promotes Baskin microflora C.C., Baskin X., Yang 165 cetfi Reports Scientific lkm .(09 vltoayAayi fMWFnto yPeoyi ecein Rescue Phenotypic by Function MBW of Analysis Evolutionary (2019) M. ulskamp ¨ 132–141. , ed:relgcldffrne rs rmvrain naaioya iechains. side arabinoxylan in variations from arise differences rheological seeds: . rnir nPatScience Plant in Frontiers 14 7 434–442. , 40672. , lkm .(09 vltoaycmaio fcmeiieprotein- competitive of comparison Evolutionary (2019) M. ulskamp ¨ 10 375. , 17 ora fEprmna Botany Experimental of Journal ln,Cl n Environment and Cell Plant, ln Physiology Plant esetvsi ln Ecology, Plant in Perspectives 178 Carbohydrate 1011–1026. , Arabi- 70 35 , , Posted on Authorea 20 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158739590.04450109 — This a preprint and has not been peer reviewed. Data may be preliminary. secretory Figure Mucilage SecretoryMucilage Cells • • • • seed: Developing • • fruit: Developing 1 Embryo Endosperm Seed coat Seed coatepidermis Released Mucilage : Fruit pericarp Fruit Fruit cells Developmental (MSCs) layer epicarp enabling epidermis mucilage scheme (dryseed dispersal) Fruit dehiscenceFruit release comparing (seed in 18 Myxospermy Dehiscent fruit - myxospermous released mucilage) the two Myxodiaspory Maturation Imbibition epidermal vs (fruit Non dehiscent fruit myxocarpous - Myxocarpy ( released mucilage) e.g. origins achene) fruit pericarp Seed coat/ of fusion species mucilage .. Posted on Authorea 20 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158739590.04450109 — This a preprint and has not been peer reviewed. Data may be preliminary. cross cross section desiccated imbibition 8 > 12 DAP> 12 mucilage development Figure - 7 DAP7 DAP6 10 DAP10 After Seed (B) (A) 2 : organization, Arabidopsis 1 Arabidopsis thaliana Arabidopsis illustrating ml2 ml1 1 and and the flax extrusion diversity as 1 model mechanisms of 1 cell species MSCprimary wall cell cell wall domain polysaccharide 19 ruptureorder amyloplasts walldomain membrane wall loosened columella mucilage mucilage primary vacuole sweling plasma cytosol layers for dynamics, upon seed water mucilage Embryo Endosperm Seed Mucilage intracellular Linum imbibition 3 ml1,ml2 ml1,ml2 ml1,ml2 Coat ml4 ml3 ml3 usitatissimum secretory secretory 2 . polysaccharidic cells cell 1 ( MSCs imbibition to mature >15DAP 12 DAP12 DAP10 (MSC) After ) Posted on Authorea 20 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158739590.04450109 — This a preprint and has not been peer reviewed. Data may be preliminary. regulator Upstream Function carbon Figure Specific Tissue Tissue / Intermediate Downstream regulators genes R2R3 MYB M complex partitioning 3 target organ master / / trait B : W The bHLH complex MYB Embryo WDR Fatty in and accumulation - bHLH bHLHs MYBs Arabidopsis toolbox Downstream acid Fatty / - endosperm GL2 WDR synthesis acid genes TTG1 TT2 (MBW) P thaliana 20 complexes Inner Proanthocyanidin seeds SK11/12 bHLHs MYBs toolbox Downstream synthesis integument Pigment Pigment through GL2 genes TTG1 regulate a P complex the Outer Mucilage MYB5 MYB61 and TT2 spatiotemporal modularity Downstream MSC MSC integument secretion genes EGL3 GL2 TT8 toolbox synthesis MYB23 TTG1 . P Posted on Authorea 20 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158739590.04450109 — This a preprint and has not been peer reviewed. Data may be preliminary. transcomplementation sequential Figure TTG1 0 EGL3* P TT8 4 : 1 Simplified evolution MYB5 TT2 2 of phylogenetic triplication and the EGL3* 3 functional A . 4 thaliana tree 5 Downstream MSC characterization MSC genes 21 of toolbox 6 land Medicago Gossypium Arabis Matthiola Arabidopsis thaliana Early Gymnosperms Amborella Zea Setaria Camellia Linum toolbox mays diverging plants usitatissimum alpina italica sinensis using incana truncatula trichopoda hirsutum studies giving land plants published . an Common R2R3 MYB 5 3 4 6 0 1 2 overview M phylogenomic Land plants Seed Flowering Eudicots Rosids Brassicaceae A. B thaliana W bHLH plants complex ancestor plants of WDR the of: , Posted on Authorea 20 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158739590.04450109 — This a preprint and has not been peer reviewed. Data may be preliminary. environmental Figure (A) Plant (A) 1 2 - - Germination control Dispersal Dispersal influence 5 : Enhancement Global development constraints overview of of . seedling seed (B) 3 - (or Water management 4 Abiotic - Soil fruit) establishment and remodeling 22 constraints mucilage Seed pericarp Fruit coat major or ecological development (C) (C) 6 - 5 Nematode - Biotic Microbial functions constraints attraction control facing