Posted on Authorea 23 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158497988.81945620 | This a preprint and has not been peer reviewed. Data may be preliminary. gao&Rlt21) hsgtee nweg e osvrlapiain npamcuia n food and extraction pharmaceutical easy 2018). in its Hoffmann as applications & well several Gaiani as Soukoulis, properties to chemical 2012; Saez- led and Amid Berger, physical knowledge & North, specific (Mirhosseini gathered mucilage 2012c; of This Huang advantage & taking Baskin 2014). industry Baskin, Ralet Yang, & 2012; Western Aguayo plant in SM (reviewed reviews studied functions recent most ecological several the 2018; addition, of Burton In characterization 2018). & structural Burton and letter & biochemical species molecular, (Phan Darwin’s model secretory the recently Charles mucilage on more the a focused of performed characterization in mainly of been morphological Review early has deep community. of scientific (MSCs) reported and the cells were composition by SM century functions, of a release analysis over ecological morphology to biochemical for its putative of and species description its 2011) of the Leubner-Metzger ability M¨uller and and (Weitbrecht, & The presence species, Its 2019). myxospermy. physical several Krause called diverse is in and & imbibition impressive Knox conferring (Galloway, upon by stems (SM) adherence secreted and mucilage or be leaves, seed can viscosity roots, mucilage fruits, as of seeds, such kinds as several such properties plants, organs called reproduction flowering of and In also range growth wide mucopolysaccharides, 2000). during a Garbary roles Anthocerotes, & protection as Nickrent dehydration such have Duff, and plants (Renzaglia, organs terrestrial around of produced are groups mucilage, diverging earliest the In the Introduction and mechanisms, molecular underlying the evolution, mucilage seed the recent These of recently support. but picture establishment roles. dissemination plant clearer ecological seed and associated a control and diversity. micro-organism germination drawing morphological soil observed to enable components. as the related such to molecular advances mostly uncovered relation its were were in of mucilage functions genes evolution exosystemic few seed sequential a some of for a carbon functions detected with balancing ecological was plants complex the polymorphism flowering Historically, high regulatory among a master level, conserved A intra-species be the diversity. and to At mechanisms natural appears molecular intra-species seed several on advances and in in recent presence inter- partitioning the Its this summarize morphological we unclear. underlying inter-species Here, are extreme functions the trait origins. whereas ecological this multiple ancestor suggests of common diversity last origins natural their evolutionary in chemical the present and was but it stages, that seedling suggests and with imbibition species early contact plant upon the flowering released in in is structures mucilage roles plant Seed play around adhesion. can present and therefore often protection features, including functions diverse numerous with providing environment, trait the plant widespread a is mucilage Polysaccharidic Abstract 2020 5, May 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 rbdpi thaliana Arabidopsis oa en&Huh 09.S ersns23 ftettlse aswt numerous with mass seed total the of 2-3% represents SM 2019). 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Data may be preliminary. io&W 07 oli,Bta,Nrh&Rlt21)o lrsrcua nlssb cnigelectron scanning by analysis ultrastructural or 2019) or Ralet seeds & hydrated North mount Botran, whole Poulain, on by 2017; obtained performed Wu are directly & generic characterizations a Qiao antibodies Further with (Ben-Tov epitope staining 2001). sections (Western section wall on or blue cell red toluidine with ruthenium as with immunolabeling such seeds stain hydrated polychromatic on (Miart and performed opening directly MSC staining pectin proper in role a (Kunieda and uncovered carefully be deposition al. all is to differential pressure in et explosion starts the polysaccharides enabling loosening simultaneously increase organized scaffold wall domain This swelling either molecular wall cell domain primary wall The occurring by 1). cell development domains polysaccharidic-proteinaceous (Figure The seed 1). wall MSCs modifications. 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(Figure penetration root and availability water period. on drying impact long major a a of following situation have SM flooding complex/natural can or more properties cycles, in physical drying role buoyancySoil and SM and wetting the run-off, investigating of water necessitate succession wind, would as with management such seeds, water imbibed in the SM and of dry (Lenser (Arshad its the dehydration respectively least for upon ways, size efficiency at dissemination and dispersal influencing structures of level their Interestingly, trade-off hydration conserve 1999b). a to its In able Gutterman to 2016). are according & 2015, imbibition change Gorb upon (Huang & can removed Kovalev (Kreitschitz, SM was properties of of mucilage Achenes dispersal properties it. their frictional on when germinate and even better adhesive and time float (Saez- longer mucilage and for un-released surface germinate with the water mutants on for or float type water can wild that provide than faster may seed SM their Saez-Aguayo imbibed Aguayo 3B-3; that mucilage, (Figure of propose lack residues a to acid have tempting galacturonic with is linkages it ionic gels, hydrophilic For highly embryo. constitute SMs As constraints the by abiotic keep fruits facing to dry Influence fruit seed-carrying the 4.2. of from dispersal 2018). it distance Erbar secreting long & by improve Fligge mucilage to (Leins, of conditions wind use windy the optimal surprising for an a dispersion. waiting reaching shows sticked, until its seeds seeds Interestingly, dispersal favors winged stops efficient 3A-2). and and an anemochory (Figure flattened by water advantage but while conditions encounters powerful hydric plant a seed for give mother the can place traits the until both conditions from of dry Combination far under dispersion dissemination seed wind-driven of prevents seeds Antitelochory on plementary. wings and SM of deserti co-occurrence The delay. without Toorop 2010; Baskin & arabicum, three namely Aethionema For myxospermy, seeds and 2018a). for dimorphic Song seeds these on & Tanveer dimorphic use species Wang, characterized same (Liu, species ving the dispersion Those and within seeds). persistence occurring non-myxospermic improve adaptation and to SM (myxospermic water contrasted morphotypes of to seed study related two gazes. the be to is to approach possibly interesting seems and Another germination water in to function permeability mucilage seed Therefore, and/or 2016). uptake Gallacher & (Geneve Santo potential (Bhatt, osmotic equal at salt amplexicaulis), with ( not species and desert application five PEG with obtained are (Geneve mucilage For without achenes explored. deeply be efficiency germination to on mucilage (Arsovski of rate effect mum2 positive germination a on suggesting delayed than also rather is germination for extrusion, observed SM rate in germination of decrease (PEG)-dependent and glycol pioneer polyethylene The Gorai Gutterman 3A-1). the in (Witztum, (Figure considering in germination time germination improve function (last also SM prevent cited supposed re-emphasized then as most regularly can and mucilage was However, excess function demonstrated. water fully This under 1969). transfer Evenari oxygen & block could of SM mucilage achene the ttg1 reii sphaerocephala Artemisia or (Gorai tal. et uat Pned20) naohrMCtobxdwsra eemtn htpeet defect a presents that mutant gene downstream toolbox MSC another On 2001). 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Gutterman & (Huang 09.Hwvr osc hntpswr bandi the in obtained were phenotypes such no However, 2009). 6 uai annua Lunaria itohru tits aslabursa-pastorisi Capsella strictus, Diptyocharpus .strictus D. .arabicum A. hc osse non-myxospermic possesses which tal. et .thaliana A. (Lu h Mtikfiesemerging fibers thick SM the , 04.Ide,mtnsthat mutants Indeed, 2014). tal. et tal. et reii sphaerocephala Artemisia Brassicaceae lpai persica Blepharis 00 and 2010) emnto deserves germination tal. et 04 huhnever though 2014) tal. et 06 allowing 2016) 07.Upon 2017). Henophyton myb61 pce ha- species the , , gl2 Posted on Authorea 23 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158497988.81945620 | This a preprint and has not been peer reviewed. Data may be preliminary. etetcudidct nivleeto Mo itccntans(iue3) novmn fS nthe on in SM shown of was Involvement plant 3C). in- (Figure the constraints metabolic for biotic this lividans community organs, on microbial plant SM of around of involvement omnipresence influence an their micro-organisms indicate and of development could importance their vestment the along consideration physiology into Taking plant environment. in the in released (Knee polysaccharides mucilage of cell border root to Similarly interactions Biotic Marsico 4.3. Di lifestyle 3B-4; non-disseminating the (Figure with soil agreement of in as environment kinds such local diverse species rheological in soil of days the improve 30 could least SM at So, for stability aggregate increase salsmn sapidmil ndwsra ee fteMCtobxcnrr oteS-eae MBW SM-related the mucilage to contrary undergoes toolbox that starting MSC pressure the species selection of between intense genes or downstream The species story. on back a mainly evolutionary characterized trace within applied mysterious intra- deeply actors is to this available the establishment molecular on The difficult these at light trait. it of shed sole both comparison makes to this variability allows on diversity great now based morphological genes shows SM toolbox astonishing that of origin This Angiosperms evolutionary levels. among the trait inter-species diverse and very species a is Myxodiaspory an PERSPECTIVES in AND development plant CONCLUSIONS the impact functions could characterized that previously functions the astonishing than new establis- manner promising seedling unexpected massive uncovered young the to be because (Roberts germination to protocarnivory level from starts nutrient development of low plant case under increases plant. a SM especially the be in hment, to for nematodes seems organism trapped it adaptation. dangerous of Surprisingly, plant death more 2018). a Provan even than & an Warren rather of (Roberts, adaptation predator nematode a the be to bursa-pastoris, could due nematode probably this more Conversely, is (Tsai attraction development. this proteins plant matodes, on and impact carbohydrates pathogenic seed-surface major have of can and plants thaliana with A. interaction close in in even are (Raviv species Nematodes independent several are of SM coat and seed secretion the protein from species acting secreted (Raviv myxospermic are stage, non old chitinase, seedling decades and the several of protease, (Hu at seeds nuclease, responsible establishment mucilage fungi as seedling with these such However, enhance interact zymes, mucilage. to significantly achene independently not by enzyme impacted probably do soil positively symbiosis on species and mycorrhizal (Hu fungal effect interaction pathogens arbuscular of positive fungal-bacterial and groups mucilage drought favor the the from to of through seedling diversity species young and same protecting the composition activities, in community explained microbial recently soil was promotion This 2012b). il- further was CO development providing hyphae plant microbial desert promoting The 3C-5) (Figure with effect” lustrated media “selective This to 2010). seeds, comparison on in co-inoculated SM in are on micro-organisms both When wilt. verticillium time longer for water retaining conductivity hydraulic from for extracted particularly SM adding modified (Deng By are 2012a). Huang properties & rheological Liu Baskin, Baskin, (Yang, mortality tal. et htihbt emnto n rwho h ahgncfungus pathogenic the of growth and germination inhibits that Mcnrbtst trc otko eaoe Fgr C6 ihteadtoa requirement additional the with 3C-6) (Figure nematodes root-knot attract to contributes SM 04.Asmlreeti rvkdby provoked is effect similar A 2014). 2 avahispanica Salvia n oul uast rmt edigetbihet(ag akn akn hn Huang & Zhang Baskin, Baskin, (Yang, establishment seedling promote to sugars soluble and lsl eaieseisof species relative closely a .thaliana A. reii sphaerocephala Artemisia ahnssativus Raphanus .dahlia V. xrtn o-deetmucilage. non-adherent excreting ceemclg and mucilage achene tal. et osdrbyrdcn h ln ies ypos(eck Schrempf & (Meschke symptoms disease plant the reducing considerably 07.Hwvr hsaiiyt ert rtisi osre nthe in conserved is proteins secrete to ability this However, 2017). .thaliana, A. ceemclg a hw ob erddb microorganisms, by degraded be to shown was mucilage achene ri the in or tal. et tal. et 01,nnahrn Mcntttsa mrsieamount impressive an constitutes SM non-adherent 2001), tal. et tal. et .hispanica S. .tain gl2 thaliana A. olttihmgraminicola Colletotrichum 7 a yopri ed loal oatatnematodes attract to able also seeds myxospermic has 09.Cnieigteprstcntr ftoene- those of nature parasitic the Considering 2019). 2017). 08.Tu,S novmn nboi interactions biotic in involvement SM Thus, 2018). .thaliana A. tal. et Madto hc ik olprilsto particles soil links which addition SM uatdpie fS ugsigthat suggesting SM of deprived mutant tal. et 09) ciesrs-soitden- stress-associated Active 2019a). etclimdahlia Verticillium .lividans S. sn h bacteria the using 09) lmrmct sone is Glomeromycota 2019b). ug (Geneve fungi a etrproliferati- better a has .brapastoris bursa C. .thaliana A. htcue the causes that Streptomyces tal. et tal. et Capsella 2018). 2017). MSC soil , Posted on Authorea 23 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158497988.81945620 — This a preprint and has not been peer reviewed. Data may be preliminary. iod .. er .. rcigo .. ebAAR lvrBJ 21)TG rtisregulate proteins TTG1 (2019) B.J. pigmentation. and Glover fate & cell A.A.R. epidermal Webb as well S.F., as Brockington activity T.J., circadian Hearn C.A., Airoldi REFERENCES https://orcid.org/0000-0003-1637-4042 Dunand Christophe of ability https://orcid.org/0000-0002-0897-6011 attraction Burlat Vincent (6) and rheologi- plant soil future (4) the ORCID as and well seed as the availability around water nematodes. establishment seed and community conferred flux microbial to water on relation anchoring of properties. in soil regulation remodelling dispersal or (3) cal seed ability through (2) sinking environment or as close species, cons- such environmental the properties facing on physical functions depending condition, major inappropriate (SM) upon mucilage seed (A) of traints. overview far Global so 3: morphology Figure SM of the actors of directs evolution Zhang downstream sequential 2019; until the members of traits in overview complex seed characterized an MBW various giving upstream of plants from specificity land toolbox spatiotemporal of (Golz the tree release regulate phylogenetic and to fied enables production modularity SM car- MYB including and spatiotemporal bHLH regulating a complexes with (MBW) in expending MYB-bHLH-WDR partitioning domains of bon wall Evolution primary of 2: pollination rupture Figure after sequential DAP=day and m4) cell. to to in (m1 cell numerous mucilage layers features from adherent the mucilage between Major different Note distinction (nam). and highly scale. layers cells four MSC mucilage to each non-adherent drawn in and domain not wall (am) are cell layers in primary features of mucilage rupture Major released multaneous the models: scale, both between similar differences at drawn are species illus- (MSC) cell del and secretory organization, mucilage imbibition. thaliana mucilage seed water Arabidopsis polysaccharidic for upon intracellular species mechanisms development, model extrusion of as diversity flax the and trating Arabidopsis 1: Figure editing. english (ANR-18-CE20- the “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 new to research uncover thankful to are track authors The promising most biotic the The environments. probably contrasted and environments. facing allowingACKNOWLEDGMENTS field particular species trait this to each diverse in related of highly functions point seed integrated studied a the SM regu- least as for this the independently functions From are times ecological Angiosperms. constraints of several across evolved range conserved have wide majoritarily a may be mucilage to complex, appears latory that complex regulatory master iu usitatissimum Linum tal. et Mmyiflec ln eeomn hog 1 oiieo eaieipc ngermination on impact negative or positive (1) through development plant influence may SM .thaliana A. 2019). ntelf aatdfo etr 01 Francoz 2001; Western from (adapted left the on .thaliana A. ntergt(dpe rmMiart from (adapted right the on (C) (Liu Mi novdi itcitrcin huh()drc ridrc influence indirect or direct (5) though interactions biotic in involved is SM tal. et ed.(A) seeds. 04 07 Li 2017; 2014, tal. et h osre neta atrrgltrTG together TTG1 regulator master ancestral conserved The 08 Li 2018; .thaliana A. 8 ieiso S eeomn ntehsoia model historical the in development MSC of Kinetics tal. et tal. et tal. et 08 Airoldi 2018; (B) rsneo ocn-hpdclmla si- columella, volcano-shaped a of presence : 09.Te30-50 The 2019). .usitatissimum L. 08 hn&Wn 2019). Wang & Chen 2018; auePlants Nature Mi oie yaitccniin in conditions abiotic by modified is SM tal. et tal. et 09,adi h mrigmo- emerging the in and 2019), 5 09 hn H & Zhang 2019; μ 1145–1153. , eunilsnhssof synthesis sequential : ieclsfo both from cells wide m .thaliana A. (B) ulskamp ¨ Simpli- MSC Posted on Authorea 23 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158497988.81945620 — This a preprint and has not been peer reviewed. Data may be preliminary. eeeRL,Hlern .. hlisTD,A-mr .&Kse ..(07 tesiflecsseed influences Stress (2017) S.T. Kester & M. Al-Amery microenvi- T.D., chia. putative mucilage-producing Phillips plants: in D.F., of germination Pec- do- Hildebrand exudates (2019) and R.L., wall V. mucilages Geneve cell Sticky value. Burlat biotechnological (2020) plant with . K. . elements . Krause remodel design & A., ronmental to P. Jauneau Knox peroxidases I., A.F., Galloway Fourquaux anchor Y., that Martinez platforms A., mains. generates Ru demethylesterification Le P., tin Ranocha Regulation cells: E., secretory Francoz mucilage seed Arabidopsis (2015) C. Dunand dynamics. & variation genes. and V. Natural Burlat underlying P., (2019) and Ranocha H.M. mucilage E., Francoz North outer seed . . in . I D., rhamnogalacturonan Poulain Sall´e for C., 181 F., in role Granier key genes A., a TTG1-like Berger reveals and G., (TTG1) Cueff 1 I., Fabrissin Glabra Testa regu- Transparent gene (2009) of V. evolution incana Hemleben The & (2019) development. A. V. trichome K. Dressel L. Gunbin thaliana & Arabidopsis D.A. controlling Afonnikov networks D.K., latory Konstantinov V., A. mucilage Doroshkov of model mathematical A of (2012) P.P.M. seeds Iannetta 419–427. & myxospermous G.R. in Squire P.E., expansion Toorop natural D.S., Jeng of W., effect Deng The (2014) P.P.M. Iannetta & P.E. conductivity. Toorop hydraulic G.R., of Squire H coatings D.S., and Jeng seed . . P.D., determination . Hallett M.C., fate W., Albani Deng 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 water S. in on biosynthesis Wang regula- anthocyanin effect & pathways mucilage S. interacting Seed Chen of (2016) desert. Arabian D. network hyper-arid Gallacher complex the & from the A. species of Santo part A., Bhatt as function, 2 COBRA-LIKE G ting 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. required M.C. secretory -L-Arabinofuranosidase McCann mucilage -D-Xylosidase/ N.C., arabidopsis Carpita bifunctional annual G.W., a the Haughn encodes T.M., in Mummenhoff Popma diaspores & A.A., dimorphic G. Arsovski of Leubner-Metzger significance V.A.A., adaptive Jansen and B., biophysics arabicum Nichols Dispersal T., Steinbrecher (2019) K., K. Sperber W., Arshad etcru scopulophilus Leptocereus Arabidopsis 1498-1518 , eeomna Cell Developmental .B.adrltdBascca n uaini h D4 motif. WD-40 the in mutation and Brassicaceae related and Br. R. (Brassicaceae). rnsi ln Science Plant in Trends edmclg oyacaiemti organization. matrix polysaccharide mucilage seed aslabursa-pastoris Capsella urn oisi eeomna Biology Developmental in Topics Current rbsalpina Arabis ln n Soil and Plant e Phytologist New 48 Ccaee ed rmPnd aazs ua. Cuba Matanzas, de Pan from seeds (Cactaceae) Arabidopsis 261-276.e8. , . ln Physiology Plant M ln Biology Plant BMC aslabursa-pastoris Capsella 387 20 rpScience Crop . .Mdk sehr’ us)o olwtrrtnin tblt and stability retention, soil-water on purse) (shepherd’s Medik. L. nentoa ora fMlclrSciences Molecular of Journal International 221 167–176. , 515–524. , ora fAi Environments Arid of Journal 1434–1446. , 9 57 150 e Phytologist New 14 2160–2169. , 1219–1234. , 131 16. , sehrspurse). (shepherds n . agB,...Hra-adS (2018) S. Harpaz-Saad . . . B., Yang M., unl ¨ 605–642. , ln Journal Plant 225 ln Biology Plant M ln Biology Plant BMC 128 1461-1469. , Botany lkm .(04 Analysis (2014) M. ulskamp ¨ 73–79. , 94 naso Botany of Annals 497–512. , 93 20 251–255. , 11 ln Physiology Plant 1–13. , 204–212. , Aethionema 19 Matthiola 53. , 109 , , Posted on Authorea 23 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158497988.81945620 — This a preprint and has not been peer reviewed. Data may be preliminary. en . lgeK ra .(08 iiu avsa al naeohr of anemochory in sails as valves Silique (2018) C. Erbar Biology & K. Fligge P., Leins in extrusion Spatiotemporal mucilage (2013) coat I. Hara-Nishimura seed 1355–1367. & for , K. required Nishitani is M., PEROXIDASE36 (2008) Nishimura of M., I. secretion Kondo Hara-Nishimura T., Embryogenesis Shimada . Regulate T., . . Kunieda Integument M., Outer Tasaka the of M., in properties Aida NARS2/NAM Arabidopsis. physical S., and in Takeda NARS1/NAC2 the M., Proteins – Ohme-Takagi Family NAC N., invasion” Mitsuda “Sticky T., (2016) Kunieda S.N. Gorb & A. lanceolata Kovalev frictio- A., and Kreitschitz adhesive Water-dependent sticking: mucilage— vs seed Slipping the Biomaterialia (2015) of S.N. of Gorb properties architecture & nal spatial A. nanoscale Kovalev and A., Kreitschitz micro- species. plant The selected (2018) of S.N. study Gorb Comparative & microstructural detailed A. A Kreitschitz mucilage? the in Root ‘stick’ source. (2001) wall wall. cell cell carbon W.D. mucilaginous 229 the coat sole Bauer does seed How the a (2017) of . . analysis as S.N. . Gorb bacteria A., & Foxworthy A. rhizosphere Kreitschitz A.R., by Jones utilization M., its Teplitski Interactions and Microbe M., rhizoids. pea and Gao from hairs F.C., mucilage root Gong of evolution E.M., The Knee (2012) L. Dolan & V.A.S. Jones China. 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 sand Cao Biophysica of desert J., mucilage interacts on Yuan the mucilage emergence Seed C., of (2019b) Wang seedling Z. D., affect Huang Huang to . . processes . R., physiochemical Liu and Z., Wang community dunes. C.C., microbial Baskin soil J.M., shrub. Baskin desert with mycorrhizal S., a Arbuscular Zhang of (2019a) D., growth Z. Hu seedling Huang & in X. functions Yang independent S., Physiology have Zhang Plant mucilage Z., Wang achene C.C., and Baskin symbiosis J.M., Baskin interactions. D., polysaccharide Hu exploiting wall cellulose: cell to and Sticking biosynthesis (2017) cellulose mucilage H.M. stand of North role & ecological J.S. the Griffiths understanding Toward shrub (2014) desert M. a stress. Neffati of osmotic of & Layers X. germination Yang (2018) seed W., M.S. in the Aloui Doblin in El production & M., mucilage A. Gorai controlling Bacic factors N.U., transcription Jayawardana of role coat. R.W., the seed Parish into S.F., Insights Li – P.J., regulation Allen J.F., Golz 9–22. , ln eladEnvironment and Cell Plant 20 ln Science Plant .se uiae. mucilage seed L. 238–243. , 17 h ln Cell Plant The ln n Soil and Plant 232 iu usitatissimum Linum 152–159. , 47 eiimperfoliatum Lepidium 14 775–787. , 1–11. , 775–784. , 272 elti ora fNanotechnology of Journal Beilstein 179–192. , 374 20 2631–2642. , 727–738. , 42 eohtndeserti Henophyton 591–605. , .se uiaiosevlp n t ilgclsignificance. biological its and envelope mucilaginous seed L. ot fia ora fBotany of Journal African South . eetana ihtpclmyxospermy. typical with annual desert A L.: LSONE PLoS lr:Mrhlg,Dsrbto,Fntoa clg fPlants of Ecology Functional Distribution, Morphology, Flora: reii sphaerocephala Artemisia 10 13 neatv ffcso eprtr,slnt and salinity temperature, of effects Interactive : e0200522.. , salJunlo ln Sciences Plant of Journal Israel 7 Arabidopsis 1918–1927. , e Phytologist New 65 naso botany of Annals Arabidopsis 187–196. , Atrca) curn nthe in occurring (Asteraceae), Lunaria edca uiaet under- to mucilage coat seed reii monosperma Artemisia (Brassicaceae). 214 caBohmc et Biochimica Acta . h ln Cell Plant The oeua Plant- Molecular 959–966. , 110 47 Arabidopsis 27–34. , 205–212. , ora of Journal Plantago Plant Acta 25 : Posted on Authorea 23 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158497988.81945620 — This a preprint and has not been peer reviewed. Data may be preliminary. agY,Wne .. ato . elGJ,WnJ,Hha . io ..(09 WD40 A (2009) R.A. Dixon . . . D., Huhman J., Wen G.J., Peel K., from Saathoff protein J.P., variants. repeat Wenger natural of Y., exploitation Pang the and for production polysaccharide perspectives 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 from F., approaches Mucilage Fournet (2018) F., M. Miart Amato . . . L., (2010) dahliae H. Cox Schrempf R., & Rossi H. V., Meschke Lanzotti R., ( (2007) chia Labella of H.M. L., fruits/seeds Scrano North . A., & mucilage Marsico A. seed Di Marion-Poll in I G., an rhamnogalacturonan Mouille of in J., potential mutation Kronenberger hydrophilic occurring O., naturally Loudet annual desert M.-C., A cold Ralet the A., in Macquet heteromorphism seed and Fruit (2010) 999–1014. C.C. , Baskin & in J.M. Advances Baskin ephemeral (2017) co-option D., and A. Tan from factor J., Gonzalez regulation. WRKY Lu protein & betalain a A. repeat for of Cantero MYB Addition WD40 A., anthocyanin model: A Bean an regulatory (2018b) A., pigment of (MBW) Campbell T. Repeat L., Xia MYB-bHLH-WD Aguirre the . A., . . Brockman A., W., Lloyd Chen X., Dai complexes. P., ternary Wang bHLH–WD40 X., Jiang sinensis H., Camellia Hou halophytes of Y., adaptation 1 Liu important GLABRA an TESTA heteromorphism: TRANSPARENT Seed (2018a) (2017) heterogeneity. J. habitat M. Song & for Chen M. Tanveer L., . Wang . . R., Liu S., Duan from and C., development Gao composition Science and C., of Plant growth Jin Variation plant D., (2016) regulates Li M.J.T. ubiquitously S., Reaney Qi K., & ( Liu 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 TESTA MYB14 and TRANSPARENT MYB5 of (2014) phosphorylation R.A. Dixon in Site-specific & (2018) J.H. Jun H. C., Yu Liu & in L. partitioning Ji carbon B., mediates Chen GLABRA1 B., Zhang C., Li in dimorphism seed and 172 fruit of plasticity and control Cevik Developmental K., Graeber T., Lenser eiaotruncatula Medicago 1691–1707. , nsesadroso rbdpi thaliana. Arabidopsis of roots and seeds on 114 10 itcoapsstrictus Diptychocarpus 684. , 1251–1263. , 254 euae nhcai n ratoyndnacmlto hog h omto fMYB– of formation the through accumulation proanthocyanidin and anthocyanin regulates 60–69. , avahispanica Salvia eiaotruncatula Medicago . ln Physiology Plant rnir nPatScience Plant in Frontiers ilg fteCell the of Biology .. Adig O.S., ¨ nentoa ora fMlclrSciences Molecular of Journal International tetmcslividans Streptomyces Bascca)adpsil dpiesignificance. adaptive possible and (Brassicaceae) . mrvssi grgt stability. aggregate soil improves L.) Arabidopsis zlN,D N., uzel ¨ 51 sncsayfrtsu-pcfi nhcai n proanthocyanidin and anthocyanin tissue-specific for necessary is 165 odRsac International Research Food Arabidopsis 2313–2326. , 1424–1439. , 111 ln n elPhysiology Cell and Plant ne .. rsh . ebe-eze .(2016) G. Leubner-Metzger . . . C., Grosche A.A., ¨ onmez ceso ffcsa affects accession irba Biotechnology Microbial 9 11 284–291. , 1515. , seeds. niistepoieaino h fungus the of proliferation the inhibits iu usitatissimum Linum Arabidopsis aueCommunications Nature h ln Cell Plant The ehoeaarabicum Aethionema β- 46 -aatsds hticessthe increases that d-galactosidase ofxalmle ( millet foxtail to ln n Soil and Plant 58 387–398. , 3 428–443. , 19 1431–1441. , 19 . cultivars. L.) E1686. , 3990–4006. , naso Botany of Annals 9 425 rnir nPlant in Frontiers 571. , . eai italica Setaria ln Physiology Plant 57–69. , International Verticillium nasof Annals 105 ). Posted on Authorea 23 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158497988.81945620 — This a preprint and has not been peer reviewed. Data may be preliminary. ukrMR,M . hnJ,NuanK,SilyNJ,Hh .. utnRA 21)Dissecting (2017) R.A. Burton . . . 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 weed A.Y.L., arable Tsai the in Co-adaptation (2012) time P.P.M. Iannetta Botany flowering & of and G.R. Squire dormancy B., seed Locardi G.S., of Begg R., Cuerva Campos P.E., Toorop 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., (2017) analysis Quian R.J. ation S., Rose Cloutier & B.J., M.L. Soto-Cerda Garg S., legume Wheeler model N., the of Approach. Smith seed X.-D., the Wang in partitioning L., He Y., polysaccharides. Song H.M. wall cell North of H.M. (2019) versatility . 1085–1116. . structure–function G.W. . North the Haughn A., demonstrates & & Berger that G.H. M.C., Dean A. Ralet K., in Marion-Poll I., Sola flotation Kronholm seed D., of A., evolution Macquet Ropartz Local C., (2014) L., Rondeau-Mouro cells. S., epidermal Botran Saez-Aguayo coat seed A., in promotes General. homogalacturonan Berger of in INHIBITOR6 methylesterification METHYLESTERASE M.-C., Myxodiaspory PECTIN on Ralet (2013) Notes with S., (Lamiaceae) Saez-Aguayo Nepetoideae Plants the of Geography in and Myxocarpy in Systematics (2001) protocarnivory facultative O. for Ryding Evidence (2018) J. Provan & pastoris J.M. Warren H.R., of Roberts innovations phylogeny. reproductive unified and Vegetative a 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 of long-term Frenkel a V., properties as Makover physicochemical functions G., and coat In Granot seed Composition L., (2019) Aghajanyan mucilage. M.C. B., seed Raviv Ralet of & of structure seeds H.M. and from North mucilage composition L., outer the Botran into D., insights Poulain New (2018) Reviews R.A. the Plant Burton Annual in & extrusion J.L. and Phan deposition mucilage for Cell required Plant ss The MYB61 (2001) S. Penfield development. trichome for not but biosynthesis 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 seeds. 109 rnir nPatScience Plant in Frontiers urn pno nFo Science Food in Opinion Current 10 13 cetfi Reports Scientific 481–489. , 1434. , 2777–2791. , 355 19 p –1 onWly&Sn,Ld hcetr UK. Chichester, Ltd, Sons, & Wiley John 1–41. pp. . 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(shepherds sn am irradiation gamma using nentoa ora of Journal International Arabidopsis 12 25 12 siliques. 99–112. , 308–323. , e0181102. , Comparative A : aslabursa- Capsella rnir in Frontiers edcoat. seed Annals 2 , Posted on Authorea 23 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158497988.81945620 — This a preprint and has not been peer reviewed. Data may be preliminary. hoX,Qa .&W ..(07 ffcieetato of extraction Effective (2017) A.M. Wu treatment. & L. Qiao X., Zhao plants. thaliana in H & 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 Yu suggests importance, T., galactoglucomannan Kotake Ecological of C.S., coating: Pereira J.J., a Lyczakowski just L., mucilage. Yu than 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 dune Baskin Degradation X., sand (2012b) Yang desert Z. a Huang of & growth W. seedling 872–883. Zhang early J.M., emergence promotes seedling Baskin improves microflora C.C., mucilage Baskin Seed (2012a) X., Z. Yang Huang shrub. & desert G. sand Liu a J.M., of Baskin C.C., during Baskin X., barrier MYB-bHLH-WDR Yang by oxygen biosynthesis flavonoid of an 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., . . Witztum Y., An J., 10.1021/acs.biomac.9b01756. of Malmstrom mucilage DOI: V.V., outer . Symonds the A.H., in structures Irani Polysaccharide V., germination Cornuault seed M.A.K., in role Williams 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 characterization in and development Isolation (2001) T.L. Western Botany Experimental of M K., Weitbrecht in G variation M.H.-W., natural Schmidt E., Zimmermann C., Voiniciuc walls. cell secondary specialized Cruciferae. produce 16 the cells G epidermal of M.H.W., coat taxonomy Schmidt the B., Yang and C., Voiniciuc structure Society Seed Linnean (1971) the J.M. of Whitehouse Journal & J.G. 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Usadel & H.M. North L., Fu M., unl ¨ 130 13 r nir nPatScience Plant in Frontiers 238–241. , Arabidopsis nentoa ora fMlclrSciences Molecular of Journal International ed iulsdb AFM. by visualised seeds ora fEprmna Botany Experimental of Journal deetse uiaeb ultrasonic by mucilage seed adherent ln,Cl n Environment and Cell Plant, ln Physiology Plant esetvsi ln Ecology, Plant in Perspectives 7 803. , Biomacromolecules 178 Arabidopsis Carbohydrate 1011–1026. , Arabidopsis Botanical Journal 70 35 seed , , Posted on Authorea 23 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158497988.81945620 — This a preprint and has not been peer reviewed. 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