Skip to main content
. 2019 Nov 28;77(11):2049–2077. doi: 10.1007/s00018-019-03388-8

Table 1.

List of the genes and mutant phenotypes described

Name Mutant Mutant phenotype Reference
AtCESA1 rsw1, any1, fxr2

Reduced cellulose content in primary cell walls, reduced growth, radial cell swelling (rsw1)

Reduced growth, swollen roots, bulging epidermal cells in cotyledons and reduced crystalline cellulose content (any1)

Resistance to flupoxam, smaller rosettes and reduced crystalline cellulose (fxr2). CESA1 null mutants are lethal

[22, 271, 272]
AtCESA2 cesa2 Phenotypically normal. CESA2 overexpression in cesa6 null mutants can rescue to some extent the cesa6-dependent growth defects [20]
AtCESA3 cev1, eli1, ixr1

Seedlings exhibit constitutive JA and ET accumulation, reductions in cellulose levels, are smaller and have roots thicker than wt (cev1)

Reduced plant growth, ectopic lignification and reduced cellulose content in seedlings (eli1)

ixr1 plants are phenotypically normal, exhibit slight reductions in crystalline cellulose levels and resistance to ISX

[2224, 26]
AtCESA4 irx5 Reduced cellulose levels in secondary cell walls. Defects in cell wall thickness in xylem vessels. Reduced mechanical strength of the stems. Adult plants are smaller than wt [273]
AtCESA5 cesa5 Mutant plants exhibit defective seed coat mucilage synthesis. CESA5 overexpression in cesa6 null mutants can rescue partially cesa6-dependent growth defects [20]
AtCESA6 prc1, ixr2

Defects in cellulose production during primary cell wall formation, short hypocotyls and ectopic lignification (prc1)

No obvious growth phenotypes but ISX resistant (ixr2)

[23, 274, 275]
AtCESA7 irx3 Reduced cellulose levels in secondary cell walls. Defects in cell wall thickness in xylem vessels. Reduced mechanical strength of the stems. Adult plants are smaller than wt [13]
AtCESA8 irx1 Reduced cellulose in secondary cell walls. Defects in cell wall thickness in xylem vessels. Reduced mechanical strength of the stems. Adult plants are smaller than wt [15]
AtCESA9 cesa9 Mutant plants display wt-like phenotype. Epidermal cells in seed coats of cesa9 seeds are distorted [276]
AtCESA10 cesa10 No obvious mutant phenotypes [17, 18]
KORRIGAN kor1 Dwarf, cellulose deficient, display defects during cytokinesis [47, 277]
KOBITO kob1 Dwarf, sterile, short hypocotyls, incomplete cell wall during cytokinesis, cellulose deficient [48]
CHITINASE-LIKE 1 ctl1/pom-pom1 Cellulose deficient, the mutation affects xyloglucan structure. Mutant plants are smaller than wt, have shorter hypocotyls and swollen roots. ctl1 phenotype is rescued by overexpressing CTL2 [49, 52]
CHITINASE-LIKE 2 ctl2 Mutant plants are similar to the wt but display ectopic lignification in stems and etiolated hypocotyls [49]
CELLULOSE SYNTHASE INTERACTING1 csi1/pom-pom2 Dwarf, short hypocotyls, swollen cells, altered microtubule organization and cellulose deficient [52, 278]
MUNC13-LIKE PROTEIN patrol1 Cellulose deficient, mutants are smaller than wt. This mutant displays short hypocotyls and roots. Adult plants are significantly smaller than the wt [32, 279]
STELLO1 stl1 No obvious mutant growth phenotype. Functionally redundant with STL2 [34]
STELLO2 stl2 No obvious mutant growth phenotype. Functionally redundant with STL1, double stl1stl2 mutants are small, display cellulose deficiency and reduced cell wall thickness [34]
 KINESIN-LIKE PROTEIN fra1 Dwarf plants with reduced mechanical strength. Mutant plants display changes in cellulose orientation with no overall reduction of cellulose [45]
CELLULOSE SYNTHASE-LIKE PROTEIN D2 csld2 Root hair formation defective with altered cytoskeleton organization [258]
CELLULOSE SYNTHASE-LIKE PROTEIN D3 csld3/kjk Root hair formation defective resulting in root hair rupture [74, 258]
CELLULOSE SYNTHASE-LIKE PROTEIN D5 csld5 No obvious growth phenotype, required for cell plate formation [75, 280]
CELLULOSE SYNTHASE-LIKE PROTEIN A7 csla7 Pollen tube growth defects and embryogenesis failure. Overexpression of CSLA9 rescues csla7 mutant phenotype [80]
CELLULOSE SYNTHASE-LIKE PROTEIN A9 csla9 Reduced glucomannan. No obvious mutant phenotypes [79, 80]
XYG-XYLOSYLTRANSFERASE xxt1 No obvious mutant phenotypes [105]
XYG-XYLOSYLTRANSFERASE xxt2 No obvious mutant growth phenotypes. Double xxt1 xxt2 show smaller rosettes and collapsed stems. Seedling hypocotyls are smaller and thicker than the wt. Cellulose deficiency associated with cellulose fibril and cortical microtubules disorganization. Lack detectable xyloglucan [105, 281]
XYLOGLUCAN GALACTOSYLTRANSFERASE  mur3-3 Cabbage-like phenotype, smaller plants with smaller rosettes. Altered xyloglucan composition. MUR3-dependent phenotype can be rescued by knocking out XXT2 alone or together with XXT1 [107, 111]
FUCOSYLTRANSFERASE 1 mur2 No obvious mutant growth phenotype, impaired in xyloglucan fucosylation [172, 282, 283]
PUTATIVE FAMILY 43 GLYCOSYL TRANSFERASE irx9 Irregular xylem phenotype, reduced cell wall thickness due to xylan chain elongation defects. Reduced xylose levels [267]
PUTATIVE FAMILY 43 GLYCOSYL TRANSFERASE irx9-like No obvious mutant phenotypes [124]
PUTATIVE FAMILY 43 GLYCOSYL TRANSFERASE irx14 Slightly smaller plants, with thinner vessels. Reduced xylose levels [123, 124]
PUTATIVE FAMILY 43 GLYCOSYL TRANSFERASE irx14-like No obvious mutant phenotypes [123]
PUTATIVE FAMILY 47 GLYCOSYL TRANSFERASE irx10 Overall normal plants with reduced xylose levels associated with a mild irregular xylem phenotype [127]
PUTATIVE FAMILY 47 GLYCOSYL TRANSFERASE irx10-like No obvious mutant phenotypes [127]
GLUCURONYL TRANSFERASE  gux1 No obvious mutant growth phenotype. Significantly reduced levels of GlcA-substituted xylan [284]
GLUCURONYL TRANSFERASE  gux2 No obvious mutant phenotype. Significantly reduced levels of GlcA-substituted xylan. Double gux1/gux2 mutants lack almost all detectable xylan substitutions [125, 284]
GLUCURONYLTRANSFERASE gux3 No obvious mutant growth phenotype. Significantly reduced levels of GlcA-substituted xylan. Triple gux1gux2 gux3 mutants are small and have reduced resistance to breaking force [133]
GLUCURONOXYLAN-METHYLTRANSERASE gxmt1-1 No obvious mutant growth phenotype. Reduced methylation of glucuronoxylan [138]
GALACTURONOSYLTRANSFERASE 1-GAUT 8 qua1/gaut8 Significantly reduced growth and decrease in both HG and xylan-synthase activity [159]
GALACTURONOSYLTRANSFERASE 10-GAUT 10 gaut10 Mutant seedlings exhibit growth defects and short roots in the absence of sucrose [162]
GALACTURONOSYLTRANSFERASE 11-GAUT 11 gaut11 Identified as HG α-GalA transferase. KO mutant seeds produce less RG I and mucilage in coat epidermal cells (SCE) [167]
PUTATIVE GLYCOSYLTRANSFERASE muci70 Required for pectin synthesis in seed coat epidermal cells. KO mutant seeds produce less mucilage [167]
PUTATIVE METHYLTRANSFERASE qua2 Defective in cell adhesion, characterized by reduced HG content [285]
PUTATIVE HOMOGALACTURONAN-METHYLTRANSFERASE qua3 No obvious growth defects and normal pectin methylation in KO plants. QUA3 RNAi suspension cells exhibit reduced pectin methylation [156]
PECTIN METHYLTRANSFERASE-CGR2 cgr2 No obvious mutant growth phenotype. Slight reduction in uronic acids and pectin methylation [161]
PECTIN METHYLTRANSFERASE- CGR3 cgr3 No obvious growth phenotypes. cgr2 cgr3 plants are dwarf, display reduced cell elongation. Double mutant plants show significant reduced degree of methylesterification, and uronic acids content compared to the single mutants, in addition to a slight cellulose reduction [161]
XYLOGALACTURONAN-XYLOSYLTRANSFERASE xgd1 No obvious mutant growth phenotype. Reduced xylose levels associated with a reduction in xylogalacturonan content [163]
GOLGI GDP-l-GALACTOSE TRANSPORTER1 gglt1/gonst3 Mutant growth phenotype is rescued by application of boric acid. Mutant lines exhibit reduced levels of L-galactose in side-chain A of RG-II [171]
GDP-MANNOSE 4,6 DEHYDRATASE ACTIVITY mur1 KO plants are defective in L-fucose, content in both pectin and hemicellulose. Reduced strength to mechanical force. Phenotype can be rescued by boric acid application [171, 173]
GALACTAN SYNTHASE 1 gals1 No obvious mutant growth phenotype. KO plants display decreased β-1,4-galactan content. Reduced galactose in stems. Decreased galactose in seeds [164]
GALACTAN SYNTHASE 2 gals2 No obvious mutant growth phenotype but decreased β-1,4-galactan content. Decreased galactose levels in seeds [164]
GALACTAN SYNTHASE 3 gals3 No obvious mutant growth phenotype. Mutant plants display decreased β-1,4-galactan content. Reduced galactose in stems [164]
RHAMNOGALACTURONAN-II XYLOSYLTRANSFERASE-1 rgxt1 No obvious mutant growth phenotype. Involved in the xylosylation of the internal fucose moiety RG II [168]
RHAMNOGALACTURONAN-II XYLOSYLTRANSFERASE-2 rgxt2 No obvious mutant growth phenotype. Involved in the xylosylation of the internal fucose moiety RG II [168]
RHAMNOGALACTURONAN-II XYLOSYLTRANSFERASE-3 rgxt4/mgp4 Short siliques. KO plants are impaired in pollen tube growth [170]
O-ACETYLTRANSFERASE  rwa2 No obvious mutant growth phenotype. KO plants characterized by reduced wall acetylation of both pectic and non-pectic polysaccharides. KO plants show increase resistance to B.cinerea [142]
O-ACETYLTRANSFERASE  tbl10 No obvious mutant growth phenotype. KO plants show reduced RG I acetylation and enhanced drought stress resistance [175]
O-ACETYLTRANSFERASE  tbl44/pmr5 KO plants show reduced rosette size. Enhanced resistance to powdery mildew and C. higginsianum. KO plants exhibit reduced pectin methyl esterification/O-acetylation associated with increased amount of pectins [151, 177]
MURUS8 mur8 KO plants have no obvious mutant growth phenotype but exhibit reductions in rhamnogalacturonan-I and rhamnose content as well as enhanced resistance to C. higginsianum [151, 179, 282]
PECTIN METHYLESTERASE 5 pme5 Involved in regulating HG methylesterification in stems. PME5 expression is reduced in the internodes of blr-6 mutants leading to reduced cell expansion and internode elongation [286]
PECTIN METHYLESTERASE INHIBITOR 3 pmei3 Involved in regulating pectin de-metylesterification in apical meristems. PMEI3 overexpression inhibits primordial formation [287]
PECTIN METHYLESTERASE 35 pme35 KO plants exhibit reduced mechanical strength in the basal part of the inflorescence stems. Reduced cell wall thickness in cortical cells of mature stems, possibly due to changes HG de-methylesterification [190]
RECEPTOR-LIKE PROTEIN 44 rlp44/cnu2 Adult KO plants exhibit stunted growth, reduced petiole length and rosette diameter. Mutant seedlings show reduced root and hypocotyl length in high sucrose levels as well as salt oversensitivity [192]
WALL-ASSOCIATED KINASE 1 wak1 No obvious mutant growth phenotype. WAK1 overexpression leads to enhance callose deposition in response to wounding and exogenous OG application. WAK1 overexpression is associated with enhanced B. cinerea resistance [288]
WALL-ASSOCIATED KINASE 2 wak2 In the absence of sucrose wak2 KO seedlings are characterized by stunted growth and short roots. Exogenous application of metabolic active sugars (fructose or glucose) helps in rescuing WAK2-dependent cell elongation defects [152]
OsCESA7 (rice) S1-24 Smaller rice plants with brittle culm phenotype. Reduced resistance to mechanical force and cellulose deficient [63]
OsCESA4 (rice) brittle culm7/bc7(t) Cellulose reduction and overall fragility of the mutant plants. Cellulose reduction is associated to enhanced lignin accumulation in stems and leaves [289]
COBRA-like protein (rice) brittle culm1 Cellulose deficient enhanced lignification and reduced resistance to mechanical force associated to brittle culm phenotype. Exhibit reduced leaf elongation [66]
COBRA-like protein—(rice) brittle culm5 Brittle culm phenotype, reduced lignification of the nodes. Cellulose and glucuronosyl arabinoxylan deficient [64]
GA20-oxidase (Sorghum) dwf1-1 Cellulose deficient mutant, dramatically reduced growth, gibberellin synthesis impaired. This mutation affects the expression of Sorghum CESA1, 3 and 6 [290]
Cellulose synthase-like F6 (rice) cslF6 Reduced plant height and smaller stems. Impaired in MLG synthesis [89]
KINESIN-LIKE PROTEIN (rice) brittle culm 12 Dramatically reduced growth, with short roots and internodes. Culms display brittle phenotype with reduced cell wall strength [68]
GLYCOSYLTRANSFERASE FAMILY 61(rice) xax1 Mutant plants exhibit reduced growth. Xylose-deficient loss-of-function rice mutant. xax1 plants are deficient in ferulic and coumaric acid [269]
PECTIN ACETYLESTERASE1 (black cottonwood) PtPAE1 Overexpression in tobacco leads to reduction in acetyl groups of pectins [182]
GALACTURONOSYLTRANSFERASE 4 (switchgrass, poplar) KD-GAUT4 (RNAi) gaut4 Involved in the synthesis of HG. Reduced expression of GAUT4 in switchgrass and poplar associated with reduced HG and RG II content. KD transgenic plants show enhanced growth when compared to wt plants [202]
POLYGALACTURONASE 1 β SUBUNIT OE:OsBURP16 Rice plants overexpressing BURP16 are phenotypically normal. At cellular level oeOsBURP16 transgenic plants show reduced cell adhesion and reduced resistance to breaking force [207]