| [1] |
Anest A, Bouchenak-Khelladi Y, Charles-Dominique T, Forest F, Caraglio Y, et al. 2024. Blocking then stinging as a case of two-step evolution of defensive cage architectures in herbivore-driven ecosystems. |
| [2] |
Satterlee JW, Alonso D, Gramazio P, Jenike KM, He J, et al. 2024. Convergent evolution of plant prickles by repeated gene co-option over deep time. |
| [3] |
Blaschek L. 2024. Well prepared: how trichome polymorphism creates an early-warning system against herbivory. |
| [4] |
Du B, Haensch R, Alfarraj S, Rennenberg H. 2024. Strategies of plants to overcome abiotic and biotic stresses. |
| [5] |
Coverdale TC. 2019. Defence emergence during early ontogeny reveals important differences between spines, thorns and prickles. |
| [6] |
Ben Lataief S, Zourgui MN, Rahmani R, Najjaa H, Gharsallah N, et al. 2021. Chemical composition, antioxidant, antimicrobial and cytotoxic activities of bioactive compounds extracted from Opuntia dillenii cladodes. |
| [7] |
Mauseth JD. 2006. Structure-function relationships in highly modified shoots of Cactaceae. |
| [8] |
Gatehouse JA. 2002. Plant resistance towards insect herbivores: a dynamic interaction. |
| [9] |
Fornara DA, Du Toit JT. 2007. Browsing lawns? Responses of Acacia nigrescens to ungulate browsing in an African savanna. |
| [10] |
Zhou N, Simonneau F, Thouroude T, Oyant LHS, Foucher F. 2021. Morphological studies of rose prickles provide new insights. |
| [11] |
Liu Y, Wang X, Li Z, Tu J, Lu YN, et al. 2023. Regulation of capsule spine formation in castor. |
| [12] |
Christin PA, Weinreich DM, Besnard G. 2010. Causes and evolutionary significance of genetic convergence. |
| [13] |
Charles-Dominique T, Davies TJ, Hempson GP, Bezeng BS, Daru BH, et al. 2016. Spiny plants, mammal browsers, and the origin of African savannas. |
| [14] |
Tomlinson KW, Yu F, Wang X, Yao X, Yu CC, et al. 2025. The macroecology of spines on woody plants. |
| [15] |
Loeuille N, Loreau M, Ferrière R. 2002. Consequences of plant-herbivore coevolution on the dynamics and functioning of ecosystems. |
| [16] |
Gong B, Zhang G. 2014. Interactions between plants and herbivores: a review of plant defense. |
| [17] |
Pei H, Wu Y, Wu W, Lyu L, Li W. 2024. A review of the types, functions and regulatory mechanisms of plant spines. |
| [18] |
Wilson-Sánchez D, Bhatia N, Runions A, Tsiantis M. 2022. From genes to shape in leaf development and evolution. |
| [19] |
de la Rosa-Manzano E, Flores J, Delgado-Sánchez P. 2016. Effects of spine-shading on aspects of photosynthesis for three cactus species. |
| [20] |
Posluszny U, Fisher JB. 2000. Thorn and hook ontogeny in Artabotrys hexapetalus (Annonaceae). |
| [21] |
Zhang Y, Zuo M, Li R, Huang J, Cheng W, et al. 2024. Morphology, structure and development of glandular prickles in the genus Rosa. |
| [22] |
Zhang F, Rossignol P, Huang T, Wang Y, May A, et al. 2020. Reprogramming of stem cell activity to convert thorns into branches. |
| [23] |
Bass E. 2025. Cutting the defense budget: how allocation costs shape induced resistance in plants. |
| [24] |
Armani M, Charles-Dominique T, Barton KE, Tomlinson KW. 2019. Developmental constraints and resource environment shape early emergence and investment in spines in saplings. |
| [25] |
Pontarp M, Petchey OL. 2018. Ecological opportunity and predator–prey interactions: linking eco-evolutionary processes and diversification in adaptive radiations. |
| [26] |
Young JP, Fulbright TE, DeYoung CA, Hewitt DG, Wester DB, et al. 2025. Shrub anti-herbivore defenses exhibit non-linear and varied responses to increased herbivore density. |
| [27] |
Strauss SY, Agrawal AA. 1999. The ecology and evolution of plant tolerance to herbivory. |
| [28] |
Leichty AR, Poethig RS. 2019. Development and evolution of age-dependent defenses in ant-acacias. |
| [29] |
Godínez-Alvarez H, Valiente-Banuet A, Rojas-Martínez A. 2002. The role of seed dispersers in the population dynamics of the columnar cactus Neobuxbaumia tetetzo. |
| [30] |
Kim ES, Mahlberg PG. 1995. Glandular cuticle formation in Cannabis (Cannabaceae). |
| [31] |
Wagner GJ. 1991. Secreting glandular trichomes: more than just hairs. |
| [32] |
Markus Lange B, Turner GW. 2013. Terpenoid biosynthesis in trichomes—current status and future opportunities. |
| [33] |
Chalvin C, Drevensek S, Dron M, Bendahmane A, Boualem A. 2020. Genetic control of glandular trichome development. |
| [34] |
Kumar P, Kumar D, Pal S, Singh S. 2025. Plant secondary metabolites in defense against phytopathogens: mechanisms, biosynthesis, and applications. |
| [35] |
Swarnkar MK, Kumar P, Dogra V, Kumar S. 2021. Prickle morphogenesis in rose is coupled with secondary metabolite accumulation and governed by canonical MBW transcriptional complex. |
| [36] |
Wu S, Song L, Chen Y, Luo C, Wan L, et al. 2026. RlTTG1 isolated from Rosa laevigata Michx. regulates trichome development and stress response in transgenic Arabidopsis. |
| [37] |
Glover BJ, Perez-Rodriguez M, Martin C. 1998. Development of several epidermal cell types can be specified by the same MYB-related plant transcription factor. |
| [38] |
Khadgi A, Weber CA. 2020. Morphological characterization of prickled and prickle-free Rubus using scanning electron microscopy. |
| [39] |
Kellogg AA, Branaman TJ, Jones NM, Little CZ, Swanson JD. 2011. Morphological studies of developing Rubus prickles suggest that they are modified glandular trichomes. |
| [40] |
Hung CY, Lin Y, Zhang M, Pollock S, Marks MD, et al. 1998. A common position-dependent mechanism controls cell-type patterning and GLABRA2 regulation in the root and hypocotyl epidermis of Arabidopsis. |
| [41] |
Pesch M, Schultheiß I, Klopffleisch K, Uhrig JF, Koegl M, et al. 2015. TRANSPARENT TESTA GLABRA1 and GLABRA1 compete for binding to GLABRA3 in Arabidopsis. |
| [42] |
Qi M, Tian X, Chen Y, Lu Y, Zhang Y. 2025. WD40 proteins PaTTG1 interact with both bHLH and MYB to regulate trichome formation and anthocyanin biosynthesis in Platanus acerifolia. |
| [43] |
Di Cristina M, Sessa G, Dolan L, Linstead P, Baima S, et al. 1996. The Arabidopsis Athb-10 (GLABRA2) is an HD-Zip protein required for regulation of root hair development. |
| [44] |
Tominaga R, Iwata M, Okada K, Wada T. 2007. Functional analysis of the epidermal-specific MYB genes CAPRICE and WEREWOLF in Arabidopsis. |
| [45] |
Gan L, Xia K, Chen JG, Wang S. 2011. Functional characterization of TRICHOMELESS2, a new single-repeat R3 MYB transcription factor in the regulation of trichome patterning in Arabidopsis. |
| [46] |
Vadde BVL, Challa KR, Sunkara P, Hegde AS, Nath U. 2019. The TCP4 transcription factor directly activates TRICHOMELESS1 and 2 and suppresses trichome initiation. |
| [47] |
Dong M, Xue S, Bartholomew ES, Zhai X, Sun L, et al. 2022. Transcriptomic and functional analysis provides molecular insights into multicellular trichome development. |
| [48] |
Su K, Sun J, Han J, Zheng T, Sun B, et al. 2022. Combined morphological and multi-omics analyses to reveal the developmental mechanism of Zanthoxylum bungeanum prickles. |
| [49] |
Chen S, Cao Y, Zhao C, Wang S, Zhang C, et al. 2026. Integrated metabolomic and transcriptomic analysis to uncover the developmental mechanism of Zanthoxylum armatum prickles. |
| [50] |
Zheng Q, Zhou S, Irish VF, Zhang F. 2026. Thorn specification in citrus plants by an SHI/STY family transcription factor. |
| [51] |
Zhu W, Zhang Z, Wu J, You Y, Bao M, et al. 2025. Transcriptome analysis reveals insights into regulatory networks of prickle formation in Rosa multiflora and the role of RmNAC43 in lignin biosynthesis during prickle hardening. |
| [52] |
Wang J, Chu Y, Yuan X, Shi X, Feng L. 2023. A CAPRICE gene of Rosa rugosa (RrCPC) suppresses the trichome formation of Arabidopsis. |
| [53] |
Huang X, Yi P, Liu Y, Li Q, Jiang Y, et al. 2022. RrTTG1 promotes fruit prickle development through an MBW complex in Rosa roxburghii. |
| [54] |
Li J, Tang B, Li Y, Li C, Guo M, et al. 2021. Rice SPL10 positively regulates trichome development through expression of HL6 and auxin-related genes. |
| [55] |
Khosla A, Paper JM, Boehler AP, Bradley AM, Neumann TR, et al. 2014. HD-zip proteins GL2 and HDG11 have redundant functions in Arabidopsis trichomes, and GL2 activates a positive feedback loop via MYB23. |
| [56] |
Wang T, Jia Q, Wang W, Hussain S, Ahmed S, et al. 2019. GCN5 modulates trichome initiation in Arabidopsis by manipulating histone acetylation of core trichome initiation regulator genes. |
| [57] |
Gan Y, Kumimoto R, Liu C, Ratcliffe O, Yu H, et al. 2006. GLABROUS INFLORESCENCE STEMS modulates the regulation by gibberellins of epidermal differentiation and shoot maturation in Arabidopsis. |
| [58] |
Zhao M, Morohashi K, Hatlestad G, Grotewold E, Lloyd A. 2008. The TTG1-bHLH-MYB complex controls trichome cell fate and patterning through direct targeting of regulatory loci. |
| [59] |
Wada T, Tachibana T, Shimura Y, Okada K. 1997. Epidermal cell differentiation in Arabidopsis determined by a Myb homolog, CPC. |
| [60] |
Schellmann S, Schnittger A, Kirik V, Wada T, Okada K, et al. 2002. TRIPTYCHON and CAPRICE mediate lateral inhibition during trichome and root hair patterning in Arabidopsis. |
| [61] |
Suo B, Seifert S, Kirik V. 2013. Arabidopsis GLASSY HAIR genes promote trichome papillae development. |
| [62] |
Zhang F, Wang Y, Irish VF. 2021. CENTRORADIALIS maintains shoot meristem indeterminacy by antagonizing THORN IDENTITY1 in Citrus. |
| [63] |
Nadakuduti SS, Pollard M, Kosma DK, Allen C Jr, Ohlrogge JB, et al. 2012. Pleiotropic phenotypes of the sticky peel mutant provide new insight into the role of CUTIN DEFICIENT2 in epidermal cell function in tomato. |
| [64] |
Ying S, Su M, Wu Y, Zhou L, Fu R, et al. 2020. Trichome regulator SlMIXTA-like directly manipulates primary metabolism in tomato fruit. |
| [65] |
Zheng F, Cui L, Li C, Xie Q, Ai G, et al. 2022. Hair interacts with SlZFP8-like to regulate the initiation and elongation of trichomes by modulating SlZFP6 expression in tomato. |
| [66] |
Zhang X, Yan F, Tang Y, Yuan Y, Deng W, et al. 2015. Auxin response gene SlARF3 plays multiple roles in tomato development and is involved in the formation of epidermal cells and trichomes. |
| [67] |
Zhang X, Chen Z, Wang C, Zhou X, Tang N, et al. 2023. Genome-wide identification of HD-ZIP gene family and screening of genes related to prickle development in Zanthoxylum armatum. |
| [68] |
Liu X, He X, Liu Z, Wu P, Tang N, et al. 2022. Transcriptome mining of genes in Zanthoxylum armatum revealed ZaMYB86 as a negative regulator of prickly development. |
| [69] |
Shan C, Dong K, Wen D, Cui Z, Cao J. 2025. A review of m6A modification in plant development and potential quality improvement. |
| [70] |
Tang N, Cao Z, Wu P, Liu Y, Lou J, et al. 2023. Comparative transcriptome analysis reveals hormone, transcriptional and epigenetic regulation involved in prickle formation in Zanthoxylum armatum. |
| [71] |
Dong Y, Li S, Wu H, Gao Y, Feng Z, et al. 2023. Advances in understanding epigenetic regulation of plant trichome development: a comprehensive review. |
| [72] |
Nyikó T, Gyula P, Ráth S, Sós-Hegedűs A, Csorba T, et al. 2025. INCREASED DNA METHYLATION 3 forms a potential chromatin remodelling complex with HAIRPLUS to regulate DNA methylation and trichome development in tomato. |
| [73] |
He L, Huang H, Bradai M, Zhao C, You Y, et al. 2022. DNA methylation-free Arabidopsis reveals crucial roles of DNA methylation in regulating gene expression and development. |
| [74] |
Patra B, Pattanaik S, Yuan L. 2013. Ubiquitin protein ligase 3 mediates the proteasomal degradation of GLABROUS 3 and ENHANCER OF GLABROUS 3, regulators of trichome development and flavonoid biosynthesis in Arabidopsis. |
| [75] |
Yu N, Cai WJ, Wang S, Shan CM, Wang LJ, et al. 2010. Temporal control of trichome distribution by microRNA156-targeted SPL genes in Arabidopsis thaliana. |
| [76] |
Vadde BVL, Challa KR, Nath U. 2018. The TCP4 transcription factor regulates trichome cell differentiation by directly activating GLABROUS INFLORESCENCE STEMS in Arabidopsis thaliana. |
| [77] |
Liu Y, Yan M, Lan H. 2026. Regulation of trichome formation by phytohormones and cytoskeleton genes in Salsola ferganica, an annual desert halophyte. |
| [78] |
Yan T, Chen M, Shen Q, Li L, Fu X, et al. 2017. HOMEODOMAIN PROTEIN 1 is required for jasmonate-mediated glandular trichome initiation in Artemisia annua. |
| [79] |
Mur LAJ, Kenton P, Atzorn R, Miersch O, Wasternack C. 2006. The outcomes of concentration-specific interactions between salicylate and jasmonate signaling include synergy, antagonism, and oxidative stress leading to cell death. |
| [80] |
Laxmi A, Paul LK, Peters JL, Khurana JP. 2004. Arabidopsis constitutive photomorphogenic mutant, bls1, displays altered brassinosteroid response and sugar sensitivity. |
| [81] |
Li QF, Wang C, Jiang L, Li S, Sun SSM, et al. 2012. An interaction between BZR1 and DELLAs mediates direct signaling crosstalk between brassinosteroids and gibberellins in Arabidopsis. |
| [82] |
Zhang L, Zhang R, Yan P, Zeng L, Zhao W, et al. 2024. PE (Prickly Eggplant) encoding a cytokinin-activating enzyme responsible for the formation of prickles in eggplant. |
| [83] |
Kumari P, Gangwar H, Gahlaut V, Kumari P, Jaiswal V. 2025. Identification of a natural RcLOG1 allele linked to prickle development in Rose (Rosa spp.). |
| [84] |
Zhou Z, Sun L, Zhao Y, An L, Yan A, et al. 2013. Zinc Finger Protein 6 (ZFP6) regulates trichome initiation by integrating gibberellin and cytokinin signaling in Arabidopsis thaliana. |
| [85] |
Plett JM, Mathur J, Regan S. 2009. Ethylene receptor ETR2 controls trichome branching by regulating microtubule assembly in Arabidopsis thaliana. |
| [86] |
Hou X, Ding L, Yu H. 2013. Crosstalk between GA and JA signaling mediates plant growth and defense. |
| [87] |
Pattanaik S, Patra B, Singh SK, Yuan L. 2014. An overview of the gene regulatory network controlling trichome development in the model plant, Arabidopsis. |
| [88] |
Tian Y, Zhao Y, Sun Y, Bo W, Huang X, et al. 2026. Genome and transcriptomics provide insights on stipular spine morphogenesis in Robinia pseudoacacia. |
| [89] |
Ren J, Duan Y, Li R, Zhang X, Shi Y, et al. 2025. Transcriptional regulation of thorn tip sclerification in plants. |
| [90] |
Endara MJ, Coley PD, Ghabash G, Nicholls JA, Dexter KG, et al. 2017. Coevolutionary arms race versus host defense chase in a tropical herbivore–plant system. |
| [91] |
Maron JL, Agrawal AA, Schemske DW. 2019. Plant–herbivore coevolution and plant speciation. |
| [92] |
Hanley ME, Lamont BB, Fairbanks MM, Rafferty CM. 2007. Plant structural traits and their role in anti-herbivore defence. |
| [93] |
Stroud JT, Ratcliff WC. 2025. Long-term studies provide unique insights into evolution. |