Comparison of Post-Germination Mobilization of Cell Wall

1981 4) L.) L.), → (1 β The patterns Glycine max ] of non-cell wall 16–18 1 − Lupin angustifolius b† Agracetus Campus, Middleton, WI and is in the public domain in the USA Lynn A Litterer The galactose mobilized from the cotyledon cell wall a,b Glycine 10–15 Correspondence to: John W Gronwald, USDA-ARS, Plant ScienceSt Research Unit, Paul, MN 55108, USA. E-mail: [email protected] Current address: Promega53711, USA. Corp., 2800 Woods HollowCurrent Rd., address: Madison, Monsanto53562, WI USA. Company, USDA-ARS, Plant Science Research Unit, St Paul, MN 55108, USA Department of Agronomy and Plant Genetics, University ofMN Minnesota, St 55108, Paul, USA The epigeal cotyledons of soybean have dual functions, serving protein content. Although the results suggest that reducing ∗ a † ‡ b makes only a small contribution to total cotyledon reserves. duced 98% and 34%, respectively, in cotyledons of seedlings galactans attached to RG-I,growth. are mobilized during early seedling roles of the high-pectinnot cell been walls defined. ofanother In legume with soybean epigeal the cotyledons, cotyledons CWP, case primarily have of lupin ( of protein, lipid and oligosaccharide mobilization have been as reserve storage and photosynthetic organs. is considered toseedling be growth. a carbon reserve that contributes to early ng early seedling growth due to mobilization and cell wall ther CWP are mobilized during early seedling growth and, if so, t cell wall polysaccharide (CWP) levels of soybean ( The lent amounts [approximately 20 mg (cotyledon pair) 4–7 ; pectin; soybean A major pectic 2 In contrast, soybean 3 Glycine max L.) seeds contain relatively Hans-Joachim G Jung, ∗ b‡ 4) galactan with a chain length The typical primary cell wall of a,b → 1,2 (1 Assuming that diverting carbon from Glycinemax β 2,8,9 : 1981–1986 www.soci.org This article is a US Government work 7 89 2009; L.) cotyledons It was postulated that synthesis of CWP in developing cell wall; cotyledon; galactose; 2,8,9

Recent research has shown that the amount of CWP in soybean dicots consists of approximately30% 35% hemicellulose pectin, on 30% a cellulose dry and weight basis. CWP synthesis duringseed cotyledon oil and/or protein, development it is would notadversely clear whether effect increase reduced seed CWP quality would or earlyserving seedling growth. a Other than compartmentation function, the structural/functional cotyledon cell walls contain approximatelycellulose 76% plus hemicellulose pectin, on and a dry 24% weight basis. seeds is negativelycontent. correlated with total seed oil and protein predominant side chain is of 43 to 47 residues. Keywords: carbohydrates (NCWC) and CWP. Galactose and arabinose14 accounted days for after 47% of planting total (DAP), CWP the ingrown levels cotyledons of under of NCWC dry a seeds. and Measured CWP 16-h weremobilized. photoperiod. re The Measured transformation 14 ofinvolving DAP, the increases cotyledon greater in uronic to than acids, a 85% glucose photosynthetic and of rhamnose. organ cotyledon was cell associatedCONCLUSION: wall with CWP galactose restructuring of plus of therestructuring arabinose soybean triggered cell was by wall light. cotyledons The amount arePublished of 2009 carbon modified by mobilized John duri Wiley & Sons, Ltd. J Sci Food Agric high levels of12% cell on wall a dry polysaccharides weight (CWP); basis. approximately The cotyledons of soybean ( INTRODUCTION Abstract BACKGROUND: In previous research, we demonstrated tha max and David A Somers carbohydrates in soybean ( John W Gronwald, of cell wall polysaccharides and non-cell wall (www.interscience.wiley.com) DOI 10.1002/jsfa.3665 Comparison of post-germination mobilization to what extent mobilization contributes to seed reserves. RESULTS: Ungerminated (dry) seeds contained equiva Short Communication Received: 13 December 2008 Revised: 22 April 2009 Accepted: 30 April 2009 Published online in Wiley Interscience: 23 June 2009 cotyledons are negatively correlated with thecotyledon sum CWP of levels seed would oil be and desirable, it is not known whe cotyledons diverts carbon fromthat protein reducing and CWP oilseed levels deposition, oil and in and/or protein. soybean cotyledons may increase polysaccharide in soybean(RG-I) cotyledons which is haspolymer rhamnogalacturonan galactose side chains and attached arabinose to homo- the rhamnose and residues. hetero- . ∗ et al )orin
: 1981–1986 89 2009; 05 level. . 05) analysis of variance . 0 0 < < P P h, (B) cell wall polysaccharides are id residues. Data points represent J Sci Food Agric between cotyledons grown under a 16-h -protected least significant difference F The treatments consisted of ungerminated 2,8,9 27 3). SE bars not shown if smaller than symbol. = n ). (A) Non-cell wall carbohydrates are defined as the sum of  SE ( Non-cell wall carbohydrates (NCWC) and cell wall polysaccharide ± darkness ( stachyose, raffinose, sucrose and starc defined as therhamnose, sum xylose, of and arabinose, totalmean fucose, uronic galactose, ac glucose, mannose, result was observed, the methodwasusedtocomparetreatmentmeans.Comparisonswere Pectin was estimated as therhamnose sum and of uronic acids arabinose, based fucose, onpectin galactose, previous composition. reports of soybean photoperiod compared to darkness at the Figure 1. (CWP) levels in cotyledons grown under a 16-h photoperiod ( indicates significant differences Statistical analysis Carbohydrate analyses weresample, with conducted each in sample consistingwere duplicate statistically of analyzed 20 on using cotyledon a completely pairs. each with randomized Data 15 design treatments. (dry) cotyledons and cotyledons collectedpost-planting, at seven grown time under intervals either lighttreatment was replicated or three times dark as described conditions. above. For Each those cotyledon traits where a significant ( For 22 Briefly, 2 . www.soci.org JW Gronwald et al Cotyledoncell This article is a US Government work and is in the public domain in the USA 2,24 15 cm (height)] containing C, photosynthetically active However, it is not known C until analyzed. × ◦ C and relative humidity was ◦ ◦ )] were harvested and the testa 80 1 , and day/night relative humidity Lipid content was calculated as − − 1 − C for 24 h prior to analysis. s 17,19–21 23 ◦ 2 Hoagland’s containing 50 ppm N. − × as described by Stombaugh (L.) Merr. cv. Lambert] seed were planted [10 25 mol m µ using galacturonic acid as a reference standard. 26 ]inpots[16cm(width) ] were weighed, placed in 50 mL conical tubes, frozen 1 1 − − Glycine max seeds (pot) Determination ofpolysaccharides soluble sugars,The defatted samples were starch analyzed for solublecellwallpolysaccharidesaspreviouslydescribed. sugars, and starch and cell wall radiation was 450 and embryonic axis were removed manually. Cotyledons(replicate) [20 pairs CWP were calculated as theglucose, mannose, sum rhamnose, xylose of and total arabinose, uronic acid residues. fucose, galactose, in liquid nitrogen and stored at Sample preparation Cotyledon samplesmeasured. were Samples were lyophilizedthree then and ground (10 s) to their burstsmill a (Foss with fine dry North powder a America,defatted using weight Eden water-cooled with Prairie, petroleum Knifetec MN, ether following USA). 1095for protocol Samples AOAC sample Soxhlet were 2003.06 lipid extraction. was 70% anddarkness, 85%, the respectively. temperature For was growth 25 under continuous the difference in dry weightsamples before were and ground after using defatting. atexture Defatted mortar and then and dried pestle at to 50 ensure a fine wallpolysaccharides(CWP)werequantifiedusingtheUppsalatotal dietary fiber method www.interscience.wiley.com/jsfa MATERIALS AND METHODS Plant material Soybean [ characterized in cotyledons of soybeanand seed during early germination seedling growth. 90%. For seedlings grown in darkness, a ‘chimney’with made bottoms from pots removed was attachedat to 4 the days upper after rim planting (DAP). ofelongating The hypocotyl the and chimney pot epicotyl. served For to both light- support andseedlings, the dark-grown pots were subirrigated asschedule required of water with or an 1/4 alternating seedlings grown in either light or dark,at cotyledons were harvested 0, 1,from 2, six pots 3, [10 seedlings 5, (pot 7, 10 and 14 DAP. At each time point, seeds starch-free, 80% ethanol-insoluble samples wereto acid hydrolyzed monosaccharide subunits. Neutral sugarsthe were alditol acetylated acetate and derivitives wereacids quantified were by quantified GC–FID. colorimetrically Uronic and by Labavitch the method of Ahmed whether structural carbon in the high-pectincotyledons cell walls of is soybean mobilizedso, during to earlyThe what seedling objectives growth extent of and,in it this if cotyledons contributes research ofphotoperiod were to soybean compared to: to seedlings total continuous (1) germinated darknessremain characterize carbon where under heterotrophic; seedlings CWP reserves. a andpatterns 16-h (2) of compare mobilization changes of(sucrose, non-cell raffinose, in stachyose, wall starch). CWP carbohydrates (NCWC) with Metro-mix200(SunGroHorticulture,Bellevue,WA,USA).Seedlings were grown in ain growth continuous chamber darkness. under For growth aday/night under 16-h temperatures the photoperiod were 16-h 25/20 or photoperiod,

1982 1983 17,20 ). (A) sucrose,  indicates significant ∗ )orindarkness(
CWP levels of seedlings grown , respectively (data not shown). www.interscience.wiley.com/jsfa 1 − 05 level. . 0 (Fig. 1). In cotyledons of seedlings grown un- 1 < − P 3). SE bars not shown if smaller than symbol. = cotyledons grown under the 16-h photoperiod orreduced in by darkness 85 was and 78%,lization in respectively. soybean This cotyledons pattern is similar of to lipid previous mobi- reports. under the 16-h photoperiod or inand darkness 53%, respectively. were reduced by 34% Non-cell wall carbohydrates Cytosolic fructose andthe glucose majority of levels samplesand were but 0.2 not when mg detected detectable (cotyledon were in pair) less than 0.5 der the 16-h photoperiod,DAP 97% (Fig. of 1A). NCWC In wereNCWC cotyledons slowed mobilized grown after by 5 in 10 days, darkness,of and NCWC after mobilization had 14 of been days, mobilized. approximately Compared 78% mobilized to during NCWC, less early CWP was seedling14 growth DAP, (Fig. the 1B). reductiongrown Between in in darkness 5 CWP compared to to levels cotyledons grownphotoperiod. was under the Measured greater 16-h 14 in DAP, cotyledons Totalnon-cellwallcarbohydratesandcellwallpolysaccharides In cotyledons ofcell ungerminated (dry) wall seeds, carbohydratessucrose) the (NCWC) levels and of (starch,similar – non- approximately cell stachyose, 11%(cotyledon of raffinose, wall pair) cotyledon dry polysaccharides weight or (CWP) 20 mg were very Approximately 85% of cotyledon sucrose wasfor mobilized by seedlings 5 grown DAP under(Fig. the 2A). 16-h Raffinose photoperiod family oraccounted oligosaccharides in (raffinose, for darkness stachyose) 36% of NCWC in ungerminated cotyledons. By n SE ( 28 ± 2cm . 0 ± oybean seedlings grown under a 16-h photoperiod ( 8 . and is in the public domain in the USA 1SE. ± : 1981–1986 This article is a US Government work 89 Cotyledon dry weight declined at an approximately 2009; 8 cm. The decline in cotyledon dry weight during 14 . Oligosaccharide and starch levels in cotyledons of s 17,20,29 0 ± 5 . Cell wall polysaccharides in soybean cotyledons www.soci.org DAP was similar for16-h seeds photoperiod germinated or and inreports. maintained darkness under a and is consistent with earlier linear rate until 10 DAP2 and weeks then under slowed either (data regime,by not cotyledon approximately shown). dry 75%. After weight Lipid was accounted reduced forweight 21% in of ungerminated cotyledon seed. dry photoperiod For or plants in grown darkness, lipid under7 declined the rapidly DAP 16-h between (data 3 not and shown). Measured 14 DAP, the lipid content of differences between cotyledons grown under a 16-h photoperiod compared to darkness at the (B) raffinose, (C) stachyose, (D) starch. Data points represent mean J Sci Food Agric Figure 2. At 14 DAP, the seedlings were inlate the leaves V1 fully stage unrolled. with When the seedlings first were trifolio- elongating grown hypocotyls in reached a darkness, maximum length of 24 at 7 DAP. Thereafter, theDAP, epicotyl–plumule the elongated length and41 at of 14 the hypocotyl plus epicotyl–plumule was Seedling growth, dry weight and lipid content Radicle emergence occurred within 24 h after planting.dons The of cotyle- seedlings germinated under continuous darkness or aphotoperiod 16-h emerged 3 to 4 DAP.16-h For photoperiod, seedlings grown the under greening the 5 cotyledons DAP. started At to 7 unfoldwith DAP, at cotyledons the fully seedlings opened were and in unifoliolate the leaves VC unrolled. stage of growth RESULTS AND DISCUSSION made for dry, ungerminated cotyledons and cotyledons harvested at each time interval, under both light andthe dark conditions, ungerminated against control cotyledonlight data. and Additionally, data dark-grown for cotyledonsinterval. Data were are presented compared as means at each time . ; 1 − et al 13–15,31 UDP-Gal : 1981–1986 34,35 89 4) galactan side The net effect of indicates significant 36 → ∗ 2009; (1 Rhamnose levels were β 4–7 respectively. Galactose does sucrose phosphate synthase (EC 32,33 J Sci Food Agric -galactosidase (EC 3.2.1.23) β 05 level. . 0 < P , Fig. 3A] is greater than the amount mobilized from raffi- 1 -galactosidase (EC 3.2.1.22), 3). SE bars not shown if smaller than symbol. − α = Compared to galactose, the arabinose level in cotyledon cell n Fig. 2B and C]. Galactose release from the and nose and stachyose reserves [3.5 mg galactose (cotyledon pair) walls is lower and mobilization occursApproximately more slowly 80% (Fig. of 3B). cell2 wall weeks arabinose of growth was in mobilized cotyledonsriod during grown or under in the 16-h darkness. photope- unchanged Cell in wall cotyledons uronic of acid seedlingsthe levels grown 14-day growth remained in period largely darkness (Fig. 3C). during However,under in cotyledons the grown 16-h photoperiod,DAP uronic were significantly acid higher levels than levels measuredgrown measured in 5–14 in cotyledons darkness. Thethe transformation increase of in the uronicAlthough cotyledon the acids to type of a occurs uronic photosynthetic during previously acid reports was organ. indicate not measured that in galacturonicnant this acid uronic study, is acid the in predomi- soybean cotyledons. these reactions is the conversion of carbonto mobilized sucrose as which galactose can be translocated to the developing seedling. not accumulate in soybeanseedling seed growth during because it germinationthe is and salvage metabolized early pathway to for nucleotide form sugar UDP-Gal synthesis via can be converted to sucrose byglucose-4-epimerase the (EC subsequent 5.1.3.2), activities of UDP- 2.4.1.14) and sucrose phosphatase (EC 3.1.3.24). chains of cotyledon cell walls andin from the raffinose cytosol oligosaccharides is catalyzed by pair) , (D) rhamnose] in cotyledon cell walls of soybean seedlings grown under SE ( ± www.soci.org JW Gronwald by 10 DAP. 1 − This article is a US Government work and is in the public domain in the USA ). Data points represent mean  oselevelsincotyledonsgrownin 4) galactan side-chains of RG-I is the → (1 β )orindarkness(
21,30 Levels of pectin sugars [(A) galactose, (B) arabinose, (C) uronic acids differences between cotyledons grown under a 16-h photoperiod compared to darkness at the www.interscience.wiley.com/jsfa Figure 3. a 16-h photoperiod ( Cell wall polysaccharides In the cotyledons ofcharide ungerminated composition seeds, was: the galactoseuronic pectic acids (48.0%), monosac- (26.0%), arabinose rhamnose (19.8%), tose (3.1%) released from and the fucose (3.1%). Galac- 3 DAP, almost(Fig. all 2B stachyose and and C).the raffinose initial In had mobilization cotyledons been of stachyose of mobilized (Fig. was 2C).Measured14DAP,stachy seedlings followed by grown an in increase darkness, darkness were 27% of levels found inseeds.Starchlevelsincotyledonsofungerminatedseedaccounted cotyledons of ungerminated for 9% of cotyledon NCWCand3DAPforcotyledonsgrownindarknessora16-hphotoperiod. (Fig. 2D). Starch increasedFor between 1 seedlings grown underdeclined the to approximately 16-h 0.4 photoperiod, mg starch (cotyledon pair) levels For cotyledons grown in darkness, slower rates ofoccurred starch after utilization starch levels peaked.starch The formation peak of in areported. transient cotyledons of germinated soybean has been predominant pectic monosaccharide mobilizedcell from walls during cotyledon 2 weeks14 of DAP, seedling approximately growth 87% (Fig. offrom 3A). cell cotyledons Measured wall grown galactose under wasness. the mobilized This 16-h is photoperiod the orin first in cotyledons report dark- of that germinated cell soybeanover a seeds. wall longer Although time galactose interval occurring is (14 days),bilized mobilized from the cotyledon amount cell of walls galactose [approximately 5.8 mo- mg (cotyledon

1984 1985 1 − ]and 10,11,14 1 − L.) where 4) galactan in → (1 β 4) galactan, is considered , respectively. Considering 1 → − ] are mobilized from cell walls Lupinus angustifolius (1 1 β − In lupin cotyledons, mobilization www.interscience.wiley.com/jsfa 12 37,38 proposed that reducing carbon flow to In large part, the greater importance of cell 2,8,9 . 12,14 In lupin cotyledons, it has been postulated that et al 16,18 and 2.2 mg (cotyledon pair) 1 − In soybean cotyledons, the amounts of galactose and arabinose For seedlings grown in darkness, the decrease in CWP during arabinose [3 mg (cotyledon pair) wall galactan as a carbonis due reserve to in the lupin significantly compared lowerAlthough lipid to mobilization content soybean of of lupin galactose cotyledons. soybean cotyledon from cell walls is notto support important seedling as growth, it a may be carbon important reserve forThe cell expansion. transformation of the cotyledonorgan to a leaf-like involves photosynthetic aexpansion. 3-fold increase in surface area due to cell of CWP, primarily galactose from to play an important roleseedling in growth. providing carbon needed to support during early seedling growth. 16-h photoperiod, theboth transformation mobilization of and the synthesis.reduction Mobilization cell of primarily cell wall wall involves galactose involves andwhich arabinose. occurs Cell upon wall cotyledon synthesis, emergence andincreased greening, results levels in of glucose, uronic acids14 and rhamnose DAP. measured The neta effect cotyledon of with mobilization less CWP and [34% synthesis less of on CWP a mg is (cotyledon pair) mobilized during earlypair) seedling growth are 5.8 mg (cotyledon 14 DAP (53% ofunder CWP) the is 16-h photoperiod greater (34% thanoccurred of in CWP). that cotyledons Cell in grown wall in cotyledons mobilization darknesswas grown over no 14 synthesis days, of but cell there wallinthelight.ThegreaterdecreaseinCWP sugars as occurs in cotyledons levelsincotyledonsgrown grown in darkness does not appear to becarbohydrate a response reserves to a to greater support need for and seedling stachyose growth. levels Lipid, measured starch 14grown DAP in are darkness higher compared in tophotoperiod. cotyledons cotyledons The reduced grown utilization under of a cytosolicincotyledonsofseedlingsgrownindarknessmayreflectadecrease 16-h carbon reserves in sink strength. After theDAP, cessation the epicotyl-plumule of elongated hypocotyl but elongation at at a slower 7 rate. Other research has shownwalls during that cell galactan expansion. decreases in plant cell CONCLUSIONS Stombaugh basis] compared to ungerminated seeds. cell wall galactan mobilizationserving has a two reserve function, functions. mobilization In maygermination be addition required cell for to post- expansion that occurs in the cotyledon. CWP (e.g. galactose, arabinose)deposition may increase in oil developing and/orthat soybean protein developing seeds. soybeancotyledons varieties Our will not with results adversely impact reduced early suggest seedlingreduced CWP growth carbon due levels reserves. to Future in research tosoybeanseedwithlowerlevelsofcotyledonaryCWPwilldetermine genetically engineer whether this strategy increases seed oil and/oridentify protein. It the may also structural/functional rolesgalactan cell of wall the in soybean high-pectin, cotyledons. high- the available reserve levels in the soybeansoluble cotyledon sugars), (lipid, the starch, carbon mobilized from themakes only cotyledon a cell minor contribution wall to total reserves. Thisto is in the contrast epigeal cotyledons of lupin ( greater amounts of galactose [19 mg (cotyledon pair) )or
3). SE bars not and is in the public domain in the USA = n SE ( ± indicates significant differences between ∗ : 1981–1986 This article is a US Government work 89 )] and the pattern of change during the 14-day 1 − 2009; ). Data points represent mean  Levels of glucose (A), xylose (B) and mannose (C) in cotyledon 05 level. . 0 Cell wall glucose levels were reduced slightly in cotyledons This study illustrates the post-germination restructuring of the < Cell wall polysaccharides in soybean cotyledons www.soci.org growth period was similar to that for rhamnose (data not shown). grown in darkness (Fig. 4A).the However, 16-h photoperiod, in cell seedlings wall growncotyledons glucose under increased are at 5 greening. DAPcontent was when detected A in cotyledons of seedlings minor grown in darkness (Fig. decline 4B). Greening of in the cotyledon was associated cell withsignificant a increase small in but wall cell wall xylose. xylose Mannose levelscell in walls cotyledon were low and exhibited agrowth gradual regime reduction (Fig. under 4C). either cell wall of soybean cotyledons. For seeds germinated under the J Sci Food Agric low in cotyledon cell wallsDAP (Fig. was 3D). associated with Greening a of slightto cotyledons increase in at cotyledons rhamnose 5 of compared seedlingscotyledon cell grown walls of in ungerminated seeds darkness.(cotyledon were Fucose also pair low levels [0.4 mg in Figure 4. cell walls of soybean seedlings grown under a 16-h photoperiod ( in darkness ( shown if smaller than symbol. cotyledons grown under a 16-h photoperiodP compared to darkness at the . in Arch Am J et al . Cucumis Am J Bot .McGraw- (L.) Merrill. 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Extraction andfrom fractionation cotyledon meal. of polysaccharides of angustifolium development. relation to seedling development andin the mobilisation of the reserves cotyledons of 160 Quantitative trait loci associated soybean seed. galactosidase or exo-(1 1 Daveby YD and Åman P, Chemical composition of certain dehulled 3 Cosgrove DJ, Assembly and enlar 2 Stombaugh SK, Jung HG, Orf JH and Somers DA, Genotypic and 8 Stombaugh SK, Orf JH, Jung HG and Somers DA, Relationships sialO otelW gnV orsnMand WhyteJNC, MorrisonIM EganSV, CottrellIW, 5 AspinallGO, 6 Brillouet J-M and Carr 7 Nakamura A, Furuta H, Maeda H, Takao T and Nagamatsu Y, Structural 4 Aspinall GO, Begbie R, Hamilton A and Whyte JNC, Polysaccharides of 9 StombaughSK,OrfJH,JungHG,ChaseK,LarkKGandSomersDA, 14 Buckeridge MS, Santos HP and Tin 15 Buckeridge MS, Hutcheon IS and Reid JSG, The role of exo-(1 16 Harris M, Mackender RO and Smith DL, Photosynthesis of cotyledons 17 Brown CS and Huber SC, Photosynthesis, reserve mobilization and 11 Parker ML, Cell wall storage polysaccharides in cotyledons of 10 Matheson NK and Saini HS, Polysaccharide and oligosaccharide 12 Crawshaw LA and Reid JSG, Changes in cell-wall polysaccharides in 13 Buckeridge MS and Reid JSG, Purification and properties of a novel www.interscience.wiley.com/jsfa REFERENCES The authors thankin Ted Jeo sample and preparationor Renee and Schirmer commercial analysis. for productspurpose assistance Mention of in of providing this traderecommendation specific publication names information or and is endorsementAgriculture. does solely by not the for imply U.S. the Department of ACKNOWLEDGEMENTS

1986