Solving the Puzzle of Shape Regulation in Plant Epidermal Pavement Cells

Literature Cited

1. 1. 

   Altartouri B, Bidhendi AJ, Tani T, Suzuki J, Conrad C et al. 2019. Pectin chemistry and cellulose crystallinity govern pavement cell morphogenesis in a multi-step mechanism. Plant Physiol 181:127–41Demonstrates that enrichment of demethylesterified homogalacturonan precedes that of cortical microtubules at periclinal wall necks during early pavement cell lobing.
   [Google Scholar]
2. 2. 

   Ambrose C, Allard JF, Cytrynbaum EN, Wasteneys GO. 2011. A CLASP-modulated cell edge barrier mechanism drives cell-wide cortical microtubule organization in Arabidopsis. Nat. Commun. 2:430
   [Google Scholar]
3. 3. 

   Ambrose JC, Shoji T, Kotzer AM, Pighin JA, Wasteneys GO. 2007. The Arabidopsis CLASP gene encodes a microtubule-associated protein involved in cell expansion and division. Plant Cell 19:2763–75
   [Google Scholar]
4. 4. 

   Amsbury S, Hunt L, Elhaddad N, Baillie A, Lundgren M et al. 2016. Stomatal function requires pectin de-methyl-esterification of the guard cell wall. Curr. Biol. 26:2899–906
   [Google Scholar]
5. 5. 

   Anderson CT, Carroll A, Akhmetova L, Somerville C. 2010. Real-time imaging of cellulose reorientation during cell wall expansion in Arabidopsis roots. Plant Physiol 152:787–96
   [Google Scholar]
6. 6. 

   Armour WJ, Barton DA, Law AM, Overall RL. 2015. Differential growth in periclinal and anticlinal walls during lobe formation in Arabidopsis cotyledon pavement cells. Plant Cell 27:2484–500
   [Google Scholar]
7. 7. 

   Askenasy E. 1870. Über den Einfluss des Wachstumsmediums auf die Gestalt der Pflanzen. Bot. Ztg. 28:193–201
   [Google Scholar]
8. 8. 

   Asl LK, Dhondt S, Boudolf V, Beemster GT, Beeckman T et al. 2011. Model-based analysis of Arabidopsis leaf epidermal cells reveals distinct division and expansion patterns for pavement and guard cells. Plant Physiol 156:2172–83
   [Google Scholar]
9. 9. 

   Augustine SM, Cherian AV, Syamaladevi DP, Subramonian N 2015. Erianthus arundinaceus HSP70 (EaHSP70) acts as a key regulator in the formation of anisotropic interdigitation in sugarcane (Saccharum spp. hybrid) in response to drought stress. Plant Cell Physiol 56:2368–80
   [Google Scholar]
10. 10. 

    Barone Lumaga M, Coiro M, Truernit E, Erdei B, De Luca P. 2015. Epidermal micromorphology in Dioon: Did volcanism constrain Dioon evolution?. Bot. J. Linnean Soc. 179:236–54
    [Google Scholar]
11. 11. 

    Baskin TI. 2005. Anisotropic expansion of the plant cell wall. Annu. Rev. Cell Dev. Biol. 21:203–22
    [Google Scholar]
12. 12. 

    Basu D, El-Assal SED, Le J, Mallery EL, Szymanski DB 2004. Interchangeable functions of Arabidopsis PIROGI and the human WAVE complex subunit SRA1 during leaf epidermal development. Development 131:4345–55
    [Google Scholar]
13. 13. 

    Basu D, Le J, El-Din El-Essal S, Huang S, Zhang C et al. 2005. DISTORTED3/SCAR2 is a putative Arabidopsis WAVE complex subunit that activates the Arp2/3 complex and is required for epidermal morphogenesis. Plant Cell 17:502–24
    [Google Scholar]
14. 14. 

    Basu D, Le J, Zakharova T, Mallery EL, Szymanski DB 2008. A SPIKE1 signaling complex controls actin-dependent cell morphogenesis through the heteromeric WAVE and ARP2/3 complexes. PNAS 105:4044–49
    [Google Scholar]
15. 15. 

    Beauzamy L, Louveaux M, Hamant O, Boudaoud A. 2015. Mechanically, the shoot apical meristem of Arabidopsis behaves like a shell inflated by a pressure of about 1 MPa. Front. Plant Sci. 6:1038
    [Google Scholar]
16. 16. 

    Bellinger MA, Sidhu SK, Rasmussen CG. 2019. Staining maize epidermal leaf peels with toluidine blue O. Bio-101e3214 https://doi.org/10.21769/BioProtoc.3214
    [Crossref] [Google Scholar]
17. 17. 

    Belteton SA, Sawchuk MG, Donohoe BS, Scarpella E, Szymanski DB. 2018. Reassessing the roles of PIN proteins and anticlinal microtubules during pavement cell morphogenesis. Plant Physiol 176:432–49
    [Google Scholar]
18. 18. 

    Bidhendi AJ, Altartouri B, Gosselin FP, Geitmann A. 2019. Mechanical stress initiates and sustains the morphogenesis of wavy leaf epidermal cells. Cell Rep 28:1237–50.e6
    [Google Scholar]
19. 19. 

    Bidhendi AJ, Geitmann A. 2018. Finite element modeling of shape changes in plant cells. Plant Physiol 176:41–56
    [Google Scholar]
20. 20. 

    Bidhendi AJ, Geitmann A. 2019. Geometrical details matter for mechanical modeling of cell morphogenesis. Dev. Cell 50:117–25.e2
    [Google Scholar]
21. 21. 

    Bou Daher F, Chen Y, Bozorg B, Clough J, Jönsson H, Braybrook SA. 2018. Anisotropic growth is achieved through the additive mechanical effect of material anisotropy and elastic asymmetry. eLife 7:e38161
    [Google Scholar]
22. 22. 

    Boudaoud A. 2010. An introduction to the mechanics of morphogenesis for plant biologists. Trends Plant Sci 15:353–60
    [Google Scholar]
23. 23. 

    Braybrook SA, Peaucelle A. 2013. Mechano-chemical aspects of organ formation in Arabidopsis thaliana: the relationship between auxin and pectin. PLOS ONE 8:e57813
    [Google Scholar]
24. 24. 

    Brembu T, Winge P, Seem M, Bones AM. 2004. NAPP and PIRP encode subunits of a putative wave regulatory protein complex involved in plant cell morphogenesis. Plant Cell 16:2335–49
    [Google Scholar]
25. 25. 

    Brenner W. 1900. Untersuchungen an einigen Fettpflanzen. Flora Oder Allg. Bot. Ztg. 87:387–439
    [Google Scholar]
26. 26. 

    Burk DH, Liu B, Zhong R, Morrison WH, Ye ZH. 2001. A katanin-like protein regulates normal cell wall biosynthesis and cell elongation. Plant Cell 13:807–27
    [Google Scholar]
27. 27. 

    Burn JE, Hocart CH, Birch RJ, Cork AC, Williamson RE. 2002. Functional analysis of the cellulose synthase genes CesA1, CesA2, and CesA3 in Arabidopsis. Plant Physiol 129:797–807
    [Google Scholar]
28. 28. 

    Burton RA, Gidley MJ, Fincher GB. 2010. Heterogeneity in the chemistry, structure and function of plant cell walls. Nat. Chem. Biol. 6:724–32
    [Google Scholar]
29. 29. 

    Cao M, Chen R, Li P, Yu Y, Zheng R et al. 2019. TMK1-mediated auxin signalling regulates differential growth of the apical hook. Nature 568:240–43
    [Google Scholar]
30. 30. 

    Chakrabortty B, Blilou I, Scheres B, Mulder BM. 2018. A computational framework for cortical microtubule dynamics in realistically shaped plant cells. PLOS Comp. Biol. 14:e1005959
    [Google Scholar]
31. 31. 

    Cosgrove DJ. 2005. Growth of the plant cell wall. Nat. Rev. Mol. Cell Biol. 6:850–61
    [Google Scholar]
32. 32. 

    Cosgrove DJ. 2016. Plant cell wall extensibility: connecting plant cell growth with cell wall structure, mechanics, and the action of wall-modifying enzymes. J. Exp. Bot. 67:463–76
    [Google Scholar]
33. 33. 

    Cosgrove DJ. 2018. Nanoscale structure, mechanics and growth of epidermal cell walls. Curr. Opin. Plant Biol. 46:77–86
    [Google Scholar]
34. 34. 

    Cosgrove DJ, Anderson CT. 2020. Plant cell growth: Do pectins drive lobe formation in Arabidopsis pavement cells?. Curr. Biol. 30:R660–62
    [Google Scholar]
35. 35. 

    Cutler SR, Ehrhardt DW, Griffitts JS, Somerville CR 2000. Random GFP::cDNA fusions enable visualization of subcellular structures in cells of Arabidopsis at a high frequency. PNAS 97:3718–23
    [Google Scholar]
36. 36. 

    Dixit R, Cyr R. 2004. Encounters between dynamic cortical microtubules promote ordering of the cortical array through angle-dependent modifications of microtubule behavior. Plant Cell 16:3274–84
    [Google Scholar]
37. 37. 

    Djakovic S, Dyachok J, Burke M, Frank MJ, Smith LG. 2006. BRICK1/HSPC300 functions with SCAR and the ARP2/3 complex to regulate epidermal cell shape in Arabidopsis. Development 133:1091–100
    [Google Scholar]
38. 38. 

    Durand-Smet P, Spelman TA, Meyerowitz EM, Jönsson H 2020. Cytoskeletal organization in isolated plant cells under geometry control. PNAS 117:17399–408
    [Google Scholar]
39. 39. 

    Elsner J, Lipowczan M, Kwiatkowska D. 2018. Differential growth of pavement cells of Arabidopsis thaliana leaf epidermis as revealed by microbead labeling. Am. J. Bot. 105:257–65
    [Google Scholar]
40. 40. 

    Elsner J, Michalski M, Kwiatkowska D. 2012. Spatiotemporal variation of leaf epidermal cell growth: a quantitative analysis of Arabidopsis thaliana wild-type and triple cyclinD3 mutant plants. Ann. Bot. 109:897–910
    [Google Scholar]
41. 41. 

    Eng RC, Sampathkumar A. 2018. Getting into shape: the mechanics behind plant morphogenesis. Curr. Opin. Plant Biol. 46:25–31
    [Google Scholar]
42. 42. 

    Feraru E, Feraru MI, Kleine-Vehn J, Martinière A, Mouille G et al. 2011. PIN polarity maintenance by the cell wall in Arabidopsis. Curr. Biol. 21:338–43
    [Google Scholar]
43. 43. 

    Frank M, Egile C, Dyachok J, Djakovic S, Nolasco M et al. 2004. Activation of Arp2/3 complex-dependent actin polymerization by plant proteins distantly related to Scar/WAVE. PNAS 101:16379–84
    [Google Scholar]
44. 44. 

    Frank MJ, Smith LG. 2002. A small, novel protein highly conserved in plants and animals promotes the polarized growth and division of maize leaf epidermal cells. Curr. Biol. 12:849–53
    [Google Scholar]
45. 45. 

    Fu Y, Gu Y, Zheng Z, Wasteneys G, Yang Z 2005. Arabidopsis interdigitating cell growth requires two antagonistic pathways with opposing action on cell morphogenesis. Cell 120:687–700Suggests that ROP-RIC signaling pathways regulate pavement cell lobe and neck formation in an antagonistic manner.
    [Google Scholar]
46. 46. 

    Fu Y, Li H, Yang Z 2002. The ROP2 GTPase controls the formation of cortical fine F-actin and the early phase of directional cell expansion during Arabidopsis organogenesis. Plant Cell 14:777–94
    [Google Scholar]
47. 47. 

    Fu Y, Xu T, Zhu L, Wen M, Yang Z 2009. A ROP GTPase signaling pathway controls cortical microtubule ordering and cell expansion in Arabidopsis. Curr. Biol. 19:1827–32
    [Google Scholar]
48. 48. 

    Fujita M, Himmelspach R, Ward J, Whittington A, Hasenbein N et al. 2013. The anisotropy1 D604N mutation in the Arabidopsis cellulose synthase1 catalytic domain reduces cell wall crystallinity and the velocity of cellulose synthase complexes. Plant Physiol 162:74–85
    [Google Scholar]
49. 49. 

    Galletti R, Ingram GC. 2015. Communication is key: Reducing DEK1 activity reveals a link between cell-cell contacts and epidermal cell differentiation status. Commun. Integr. Biol. 8:e1059979
    [Google Scholar]
50. 50. 

    Gao Y, Zhang Y, Zhang D, Dai X, Estelle M, Zhao Y 2015. Auxin binding protein 1 (ABP1) is not required for either auxin signaling or Arabidopsis development. PNAS 112:2275–80
    [Google Scholar]
51. 51. 

    Geisler M, Nadeau J, Sack FD. 2000. Oriented asymmetric divisions that generate the stomatal spacing pattern in Arabidopsis are disrupted by the too many mouths mutation. Plant Cell 12:2075–86
    [Google Scholar]
52. 52. 

    Glover BJ. 2000. Differentiation in plant epidermal cells. J. Exp. Bot. 51:497–505
    [Google Scholar]
53. 53. 

    Gonzalez N, Vanhaeren H, Inzé D. 2012. Leaf size control: complex coordination of cell division and expansion. Trends Plant Sci 17:332–40
    [Google Scholar]
54. 54. 

    Grones P, Chen X, Simon S, Kaufmann WA, De Rycke R et al. 2015. Auxin-binding pocket of ABP1 is crucial for its gain-of-function cellular and developmental roles. J. Exp. Bot. 66:5055–65
    [Google Scholar]
55. 55. 

    Grones P, Majda M, Doyle SM, Van Damme D, Robert S 2020. Fluctuating auxin response gradients determine pavement cell-shape acquisition. PNAS 117:16027–34Reveals the importance of fluctuating auxin response gradients for lobe formation in pavement cells within spiral stomatal complexes.
    [Google Scholar]
56. 56. 

    Guimil S, Dunand C. 2007. Cell growth and differentiation in Arabidopsis epidermal cells. J. Exp. Bot. 58:3829–40
    [Google Scholar]
57. 57. 

    Guo X, Qin Q, Yan J, Niu Y, Huang B et al. 2015. TYPE-ONE PROTEIN PHOSPHATASE4 regulates pavement cell interdigitation by modulating PIN-FORMED1 polarity and trafficking in Arabidopsis. Plant Physiol 167:1058–75
    [Google Scholar]
58. 58. 

    Haas KT, Wightman R, Meyerowitz EM, Peaucelle A. 2020. Pectin homogalacturonan nanofilament expansion drives morphogenesis in plant epidermal cells. Science 367:1003–7Presents a new theory of demethylesterification-driven expansion of local anticlinal homogalacturonan nanofilaments inducing pavement cell lobing, supported by computational modeling.
    [Google Scholar]
59. 59. 

    Hamant O, Heisler MG, Jönsson H, Krupinski P, Uyttewaal M et al. 2008. Developmental patterning by mechanical signals in Arabidopsis. Science 322:1650–55
    [Google Scholar]
60. 60. 

    Hamant O, Inoue D, Bouchez D, Dumais J, Mjolsness E. 2019. Are microtubules tension sensors?. Nat. Commun. 10:2360
    [Google Scholar]
61. 61. 

    Hazak O, Bloch D, Poraty L, Sternberg H, Zhang J et al. 2010. A rho scaffold integrates the secretory system with feedback mechanisms in regulation of auxin distribution. PLOS Biol 8:e1000282
    [Google Scholar]
62. 62. 

    Hazak O, Obolski U, Prat T, Friml J, Hadany L, Yalovsky S 2014. Bimodal regulation of ICR1 levels generates self-organizing auxin distribution. PNAS 111:E5471–79
    [Google Scholar]
63. 63. 

    Hejnowicz Z, Rusin A, Rusin T. 2000. Tensile tissue stress affects the orientation of cortical microtubules in the epidermis of sunflower hypocotyl. J. Plant Growth Regul. 19:31–44
    [Google Scholar]
64. 64. 

    Hervieux N, Dumond M, Sapala A, Routier-Kierzkowska AL, Kierzkowski D et al. 2016. A mechanical feedback restricts sepal growth and shape in Arabidopsis. Curr. Biol 26:1019–28
    [Google Scholar]
65. 65. 

    Higaki T, Kutsuna N, Akita K, Takigawa-Imamura H, Yoshimura K, Miura T. 2016. A theoretical model of jigsaw-puzzle pattern formation by plant leaf epidermal cells. PLOS Comput. Biol. 12:e1004833
    [Google Scholar]
66. 66. 

    Higaki T, Takigawa-Imamura H, Akita K, Kutsuna N, Kobayashi R et al. 2017. Exogenous cellulase switches cell interdigitation to cell elongation in an RIC1-dependent manner in Arabidopsis thaliana cotyledon pavement cells. Plant Cell Physiol 58:106–19
    [Google Scholar]
67. 67. 

    Ivakov A, Persson S. 2013. Plant cell shape: modulators and measurements. Front. Plant Sci. 4:439
    [Google Scholar]
68. 68. 

    Jacques E, Verbelen J-P, Vissenberg K. 2013. Mechanical stress in Arabidopsis leaves orients microtubules in a ‘continuous’ supracellular pattern. BMC Plant Biol 13:163
    [Google Scholar]
69. 69. 

    Jacques E, Verbelen J-P, Vissenberg K. 2014. Review on shape formation in epidermal pavement cells of the Arabidopsis leaf. Funct. Plant Biol. 41:914–21
    [Google Scholar]
70. 70. 

    Javelle M, Vernoud V, Rogowsky PM, Ingram GC. 2011. Epidermis: the formation and functions of a fundamental plant tissue. New Phytol 189:17–39
    [Google Scholar]
71. 71. 

    Jooste M, Dreyer LL, Oberlander KC. 2016. The phylogenetic significance of leaf anatomical traits of southern African Oxalis. BMC Evol. Biol. 16:225
    [Google Scholar]
72. 72. 

    Kerstens S, Decraemer WF, Verbelen JP. 2001. Cell walls at the plant surface behave mechanically like fiber-reinforced composite materials. Plant Physiol 127:381–85
    [Google Scholar]
73. 73. 

    Landrein B, Hamant O. 2013. How mechanical stress controls microtubule behavior and morphogenesis in plants: history, experiments and revisited theories. Plant J 75:324–38
    [Google Scholar]
74. 74. 

    Lavy M, Bloch D, Hazak O, Gutman I, Poraty L et al. 2007. A novel ROP/RAC effector links cell polarity, root-meristem maintenance, and vesicle trafficking. Curr. Biol. 17:947–52
    [Google Scholar]
75. 75. 

    Le J, El-Assal SED, Basu D, Saad ME, Szymanski DB 2003. Requirements for Arabidopsis ATARP2 and ATARP3 during epidermal development. Curr. Biol. 13:1341–47
    [Google Scholar]
76. 76. 

    Le J, Liu XG, Yang KZ, Chen XL, Zou JJ et al. 2014. Auxin transport and activity regulate stomatal patterning and development. Nat. Commun. 5:3090
    [Google Scholar]
77. 77. 

    Li H, Lin D, Dhonukshe P, Nagawa S, Chen D et al. 2011. Phosphorylation switch modulates the interdigitated pattern of PIN1 localization and cell expansion in Arabidopsis leaf epidermis. Cell Res 21:970–78
    [Google Scholar]
78. 78. 

    Li H, Xu T, Lin D, Wen M, Xie M et al. 2013. Cytokinin signaling regulates pavement cell morphogenesis in Arabidopsis. Cell Res 23:290–99
    [Google Scholar]
79. 79. 

    Li J, Kim T, Szymanski DB. 2019. Multi-scale regulation of cell branching: modeling morphogenesis. Dev. Biol. 451:40–52
    [Google Scholar]
80. 80. 

    Li S, Blanchoin L, Yang Z, Lord EM 2003. The putative Arabidopsis Arp2/3 complex controls leaf cell morphogenesis. Plant Physiol 132:2034–44
    [Google Scholar]
81. 81. 

    Lin D, Cao L, Zhou Z, Zhu L, Ehrhardt D et al. 2013. Rho GTPase signaling activates microtubule severing to promote microtubule ordering in Arabidopsis. Curr. Biol. 23:290–97
    [Google Scholar]
82. 82. 

    Lin D, Ren H, Fu Y. 2015. ROP GTPase-mediated auxin signaling regulates pavement cell interdigitation in Arabidopsis thaliana. J. Integr. Plant Biol. 57:31–39
    [Google Scholar]
83. 83. 

    Liu X, Yang Q, Wang Y, Wang L, Fu Y, Wang X. 2018. Brassinosteroids regulate pavement cell growth by mediating BIN2-induced microtubule stabilization. J. Exp. Bot. 69:1037–49
    [Google Scholar]
84. 84. 

    Majda M, Grones P, Sintorn IM, Vain T, Milani P et al. 2017. Mechanochemical polarization of contiguous cell walls shapes plant pavement cells. Dev. Cell 43:290–304.e4Suggests roles for anticlinal wall mechanical and chemical gradients in pavement cell interdigitation, supported by a finite-element-model-based simulation.
    [Google Scholar]
85. 85. 

    Majda M, Krupinski P, Jönsson H, Hamant O, Robert S 2019. Mechanical asymmetry of the cell wall predicts changes in pavement cell geometry. Dev. Cell 50:9–10
    [Google Scholar]
86. 86. 

    Majda M, Robert S 2018. The role of auxin in cell wall expansion. Int. J. Mol. Sci. 19:951
    [Google Scholar]
87. 87. 

    Malinowski R. 2013. Understanding of leaf development—the science of complexity. Plants 2:396–415
    [Google Scholar]
88. 88. 

    Mathur J. 2004. Cell shape development in plants. Trends Plant Sci 9:583–90
    [Google Scholar]
89. 89. 

    Mathur J, Mathur N, Kernebeck B, Hülskamp M. 2003. Mutations in actin-related proteins 2 and 3 affect cell shape development in Arabidopsis. Plant Cell 15:1632–45
    [Google Scholar]
90. 90. 

    McKenna JF, Rolfe DJ, Webb SED, Tolmie AF, Botchway SW et al. 2019. The cell wall regulates dynamics and size of plasma-membrane nanodomains in Arabidopsis. PNAS 116:12857–62
    [Google Scholar]
91. 91. 

    Mirabet V, Krupinski P, Hamant O, Meyerowitz EM, Jönsson H, Boudaoud A. 2018. The self-organization of plant microtubules inside the cell volume yields their cortical localization, stable alignment, and sensitivity to external cues. PLOS Comp. Biol. 14:e1006011
    [Google Scholar]
92. 92. 

    Nagawa S, Xu T, Lin D, Dhonukshe P, Zhang X et al. 2012. ROP GTPase-dependent actin microfilaments promote PIN1 polarization by localized inhibition of clathrin-dependent endocytosis. PLOS Biol 10:e1001299
    [Google Scholar]
93. 93. 

    Onoda Y, Schieving F, Anten NP. 2015. A novel method of measuring leaf epidermis and mesophyll stiffness shows the ubiquitous nature of the sandwich structure of leaf laminas in broad-leaved angiosperm species. J. Exp. Bot. 66:2487–99
    [Google Scholar]
94. 94. 

    Paciorek T, Zažímalová E, Ruthardt N, Petrášek J, Stierhof Y-D et al. 2005. Auxin inhibits endocytosis and promotes its own efflux from cells. Nature 435:1251–56
    [Google Scholar]
95. 95. 

    Pan X, Fang L, Liu J, Senay-Aras B, Lin W et al. 2020. Auxin-induced signal protein nanoclustering contributes to cell polarity formation. Nat. Commun. 11:3914Demonstrates that auxin reorganizes cortical microtubules to establish pavement cell polarity by promoting TMK1 and ROP6 nanocluster formation across the plasma membrane.
    [Google Scholar]
96. 96. 

    Panikashvili D, Shi JX, Schreiber L, Aharoni A. 2009. The Arabidopsis DCR encoding a soluble BAHD acyltransferase is required for cutin polyester formation and seed hydration properties. Plant Physiol 151:1773–89
    [Google Scholar]
97. 97. 

    Panteris E, Apostolakos P, Galatis B. 1993. Microtubule organization and cell morphogenesis in two semi-lobed cell types of Adiantum capillus-veneris L. leaflets. New Phytol 125:509–20
    [Google Scholar]
98. 98. 

    Panteris E, Apostolakos P, Galatis B. 1994. Sinuous ordinary epidermal cells: behind several patterns of waviness, a common morphogenetic mechanism. New Phytol 127:771–80Reveals the presence of cortical microtubule arrays at anticlinal and periclinal wall neck regions in lobed pavement cells of several species.
    [Google Scholar]
99. 99. 

    Panteris E, Galatis B. 2005. The morphogenesis of lobed plant cells in the mesophyll and epidermis: organization and distinct roles of cortical microtubules and actin filaments. New Phytol 167:721–32
    [Google Scholar]
100. 100. 

     Paredez AR, Somerville CR, Ehrhardt DW. 2006. Visualization of cellulose synthase demonstrates functional association with microtubules. Science 312:1491–95
     [Google Scholar]
101. 101. 

     Park YB, Cosgrove DJ. 2015. Xyloglucan and its interactions with other components of the growing cell wall. Plant Cell Physiol 56:180–94
     [Google Scholar]
102. 102. 

     Parre E, Geitmann A. 2005. Pectin and the role of the physical properties of the cell wall in pollen tube growth of Solanum chacoense. Planta 220:582–92
     [Google Scholar]
103. 103. 

     Peaucelle A, Braybrook SA, Le Guillou L, Bron E, Kuhlemeier C, Höfte H 2011. Pectin-induced changes in cell wall mechanics underlie organ initiation in Arabidopsis. Curr. Biol. 21:1720–26
     [Google Scholar]
104. 104. 

     Peaucelle A, Wightman R, Höfte H. 2015. The control of growth symmetry breaking in the Arabidopsis hypocotyl. Curr. Biol. 25:1746–52
     [Google Scholar]
105. 105. 

     Pigg K. 1990. Anatomically preserved Glossopteris foliage from the central Transantarctic Mountains. Rev. Palaeobot. Palynol. 66:105–27
     [Google Scholar]
106. 106. 

     Pillitteri LJ, Dong J 2013. Stomatal development in Arabidopsis. Arabidopsis Book 11:e0162
     [Google Scholar]
107. 107. 

     Popper ZA, Michel G, Hervé C, Domozych DS, Willats WG et al. 2011. Evolution and diversity of plant cell walls: from algae to flowering plants. Annu. Rev. Plant Biol. 62:567–90
     [Google Scholar]
108. 108. 

     Poulson M, Vogelmann T. 1990. Epidermal focussing and effects upon photosynthetic light-harvesting in leaves of Oxalis. Plant Cell Environ 13:803–11
     [Google Scholar]
109. 109. 

     Pyke K, Marrison J, Leech R. 1991. Temporal and spatial development of the cells of the expanding first leaf of Arabidopsis thaliana (L.) Heynh. J. Exp. Bot. 42:1407–16
     [Google Scholar]
110. 110. 

     Qiu JL, Jilk R, Marks MD, Szymanski DB. 2002. The Arabidopsis SPIKE1 gene is required for normal cell shape control and tissue development. Plant Cell 14:101–18
     [Google Scholar]
111. 111. 

     Ren H, Dang X, Yang Y, Huang D, Liu M et al. 2016. SPIKE1 activates ROP GTPase to modulate petal growth and shape. Plant Phys 172:358–71
     [Google Scholar]
112. 112. 

     Robert S, Kleine-Vehn J, Barbez E, Sauer M, Paciorek T et al. 2010. ABP1 mediates auxin inhibition of clathrin-dependent endocytosis in Arabidopsis. Cell 143:111–21
     [Google Scholar]
113. 113. 

     Rongpipi S, Ye D, Gomez ED, Gomez EW. 2019. Progress and opportunities in the characterization of cellulose—an important regulator of cell wall growth and mechanics. Front. Plant Sci. 9:1894
     [Google Scholar]
114. 114. 

     Sampathkumar A, Krupinski P, Wightman R, Milani P, Berquand A et al. 2014. Subcellular and supracellular mechanical stress prescribes cytoskeleton behavior in Arabidopsis cotyledon pavement cells. eLife 3:e01967Reveals periclinal neck stiffness patterns matching cortical microtubule arrays and shows microtubule alignment with predicted stress patterns under compression, in pavement cells.
     [Google Scholar]
115. 115. 

     Sampathkumar A, Yan A, Krupinski P, Meyerowitz EM. 2014. Physical forces regulate plant development and morphogenesis. Curr. Biol. 24:R475–83
     [Google Scholar]
116. 116. 

     Samuels L, Kunst L, Jetter R. 2008. Sealing plant surfaces: cuticular wax formation by epidermal cells. Annu. Rev. Plant Biol. 59:683–707
     [Google Scholar]
117. 117. 

     Sapala A, Runions A, Routier-Kierzkowska AL, Das Gupta M, Hong L et al. 2018. Why plants make puzzle cells, and how their shape emerges. eLife 7:e32794Proposes that cell interdigitation serves to reduce cell wall mechanical stress in the epidermis of isotropically growing organs.
     [Google Scholar]
118. 118. 

     Sapala A, Runions A, Smith RS. 2019. Mechanics, geometry and genetics of epidermal cell shape regulation: different pieces of the same puzzle. Curr. Opin. Plant Biol. 47:1–8
     [Google Scholar]
119. 119. 

     Schnurr J, Shockey J, Browse J. 2004. The Acyl-CoA synthetase encoded by LACS2 is essential for normal cuticle development in Arabidopsis. Plant Cell 16:629–42
     [Google Scholar]
120. 120. 

     Shpak ED, McAbee JM, Pillitteri LJ, Torii KU. 2005. Stomatal patterning and differentiation by synergistic interactions of receptor kinases. Science 309:290–93
     [Google Scholar]
121. 121. 

     Skalák J, Vercruyssen L, Claeys H, Hradilová J, Černý M et al. 2019. Multifaceted activity of cytokinin in leaf development shapes its size and structure in Arabidopsis. Plant J 97:805–24
     [Google Scholar]
122. 122. 

     Smith LG. 2003. Cytoskeletal control of plant cell shape: getting the fine points. Curr. Opin. Plant Biol. 6:63–73
     [Google Scholar]
123. 123. 

     Sotiriou P, Giannoutsou E, Panteris E, Galatis B, Apostolakos P. 2018. Local differentiation of cell wall matrix polysaccharides in sinuous pavement cells: its possible involvement in the flexibility of cell shape. Plant Biol 20:223–37
     [Google Scholar]
124. 124. 

     Staff L, Hurd P, Reale L, Seoighe C, Rockwood A, Gehring C. 2012. The hidden geometries of the Arabidopsis thaliana epidermis. PLOS ONE 7:e43546
     [Google Scholar]
125. 125. 

     Szymanski DB. 2014. The kinematics and mechanics of leaf expansion: new pieces to the Arabidopsis puzzle. Curr. Opin. Plant Biol. 22:141–48
     [Google Scholar]
126. 126. 

     Tao LZ, Cheung AY, Wu HM. 2002. Plant Rac-like GTPases are activated by auxin and mediate auxin-responsive gene expression. Plant Cell 14:2745–60
     [Google Scholar]
127. 127. 

     Vőfély RV, Gallagher J, Pisano GD, Bartlett M, Braybrook SA. 2019. Of puzzles and pavements: a quantitative exploration of leaf epidermal cell shape. New Phytol 221:540–52Presents extensive quantitative analyses of pavement cell shape at phylogenetic scales and suggests that no single common reason underlies puzzle shape evolution.
     [Google Scholar]
128. 128. 

     Vogelmann TC, Bornman JF, Yates DJ. 1996. Focusing of light by leaf epidermal cells. Physiol. Plant 98:43–56
     [Google Scholar]
129. 129. 

     Wang G, Wang C, Liu W, Ma Y, Dong L et al. 2018. Augmin antagonizes katanin at microtubule crossovers to control the dynamic organization of plant cortical arrays. Curr. Biol. 28:1311–17.e3
     [Google Scholar]
130. 130. 

     Watson R. 1942. The effect of cuticular hardening on the form of epidermal cells. New Phytol 41:223–29
     [Google Scholar]
131. 131. 

     Wightman R, Chomicki G, Kumar M, Carr P, Turner SR. 2013. SPIRAL2 determines plant microtubule organization by modulating microtubule severing. Curr. Biol. 23:1902–7
     [Google Scholar]
132. 132. 

     Wilmoth JC, Wang S, Tiwari SB, Joshi AD, Hagen G et al. 2005. NPH4/ARF7 and ARF19 promote leaf expansion and auxin-induced lateral root formation. Plant J 43:118–30
     [Google Scholar]
133. 133. 

     Wong JH, Kato T, Belteton SA, Shimizu R, Kinoshita N et al. 2019. Basic proline-rich protein-mediated microtubules are essential for lobe growth and flattened cell geometry. Plant Physiol 181:1535–51
     [Google Scholar]
134. 134. 

     Xu T, Dai N, Chen J, Nagawa S, Cao M et al. 2014. Cell surface ABP1-TMK auxin-sensing complex activates ROP GTPase signaling. Science 343:1025–28Shows that auxin regulates pavement cell shape via TMK1-mediated activation of ROPs at the plasma membrane.
     [Google Scholar]
135. 135. 

     Xu T, Wen M, Nagawa S, Fu Y, Chen J-G et al. 2010. Cell surface- and Rho GTPase-based auxin signaling controls cellular interdigitation in Arabidopsis. Cell 143:99–110
     [Google Scholar]
136. 136. 

     Zhang C, Halsey LE, Szymanski DB. 2011. The development and geometry of shape change in Arabidopsis thaliana cotyledon pavement cells. BMC Plant Biol 11:27
     [Google Scholar]
137. 137. 

     Zhang C, Kotchoni SO, Samuels AL, Szymanski DB. 2010. SPIKE1 signals originate from and assemble specialized domains of the endoplasmic reticulum. Curr. Biol. 20:2144–49
     [Google Scholar]
138. 138. 

     Zhang C, Mallery E, Reagan S, Boyko VP, Kotchoni SO, Szymanski DB. 2013. The endoplasmic reticulum is a reservoir for WAVE/SCAR regulatory complex signaling in the Arabidopsis leaf. Plant Physiol 162:689–706
     [Google Scholar]
139. 139. 

     Zhang C, Mallery EL, Schlueter J, Huang S, Fan Y et al. 2008. Arabidopsis SCARs function interchangeably to meet actin-related protein 2/3 activation thresholds during morphogenesis. Plant Cell 20:995–1011
     [Google Scholar]
140. 140. 

     Zhang C, Mallery EL, Szymanski DB. 2013. ARP2/3 localization in Arabidopsis leaf pavement cells: a diversity of intracellular pools and cytoskeletal interactions. Front. Plant Sci. 4:238
     [Google Scholar]
141. 141. 

     Zhou W, Wang Y, Wu Z, Luo L, Liu P et al. 2016. Homologs of SCAR/WAVE complex components are required for epidermal cell morphogenesis in rice. J. Exp. Bot. 67:4311–23
     [Google Scholar]
142. 142. 

     Zienkiewicz OC, Taylor RL. 2005. The Finite Element Method for Solid and Structural Mechanics Oxford, UK: Elsevier