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, Integration of the environmental signals is 38 achieved by a number of growth-promoting and growth-restricting factors. Many 39 genes with growth-promoting and growth-restricting functions that act on cell 40 division or on cell expansion have been identified in plants and in animals (B? ogre 41 et al, Developmental processes largely depend on both genetic factors and environ34 mental inputs in animals and plants, vol.37, 2006.
, However, these genes act in independent genetic pathways, 44 making it difficult to develop an integrated and comprehensive model of organ 45 size control. 46 A common pathway controlling animal as well as plant organ growth has been 47 proposed to involve the Translationally Controlled Tumor Protein (TCTP), 2011.
, TCTP is conserved among all eukaryotes and has been proposed to 51 have diverse roles in developmental and defense processes, including several 52 cellular functions such as cell proliferation, expansion, and death (Bommer and 53, Thiele, 2004.
, we will present and discuss the up-to-date knowledge on the 55 multiple roles of TCTP in growth and in response to various signals and stresses in 56 plants, and how gained knowledge help understanding the TCTP's central role in 57 plant and animal development
, Since the early 1990s, homologs of TCTP have been identified in many plant 60 species, such as the model plant Arabidopsis thaliana, alfalfa, pea, Pharbitis nil, 61 grape vine, oil palm, Jatropha curcas, cassava, cabbage, rice, strawberry, tobacco, 62 rubber tree, 1992.
,
, Lopez and Franco, vol.64, 2003.
, TCTP is generally present as a single gene copy in the genome, but 67 many species carry more than one gene, 2014.
, Mammals seem to have many TCTP gene copies in their 69 genome, but only one is likely functional, 2000.
, Plant species harbor one and up to five TCTP gene copies
, Three distinct structural domains are found in TCTP: a core ?-sheet 134 domain, an ?-helical domain, and a flexible loop structure. Major differences 135 between plant and animal TCTPs are observed in the flexible loop, but many of these gene copies 132 human structures, 2005.
, Berkowitz et al, 2008.
, In plants, the 139 predictive structures of TCTP support the phylogenetic evidence that they fall 140 into two sub-clades, AtTCTP1-like and CmTCTP-like (Cucurbita max.), that differ 141 in the structure of the central "pocket" region and in the flexible loop, suggesting 142 different functions. It should be noted that in plant species harboring a single TCTP 143 gene, the sequence is usually AtTCTP1-like, 2008.
, Considering the high degree of amino-acids conservation and the high similarity 145 of predictive tri-dimensional structure among phyla, it is tempting to suggest that 146 plant and animal TCTPs share many of their roles. In agreement with this hypoth147 esis, Brioudes et al. (2010) demonstrated in vivo that Drosophila dTCTP could 148 complement cell proliferation defects associated with AtTCTP loss-of-function in 149 Arabidopsis and vice versa
, One could imagine that despite 157 the obvious absence of primary amino acid similarities of TCTP interacting pro158 teins, other functionally conserved proteins can fulfill their role and secure TCTP 159 signaling in plants. Such information might give precious indications about evolu160 tion of TCTP function and interactors across kingdoms. in plants as in animals. Final organ shape and size are the result of 164 coordinated cell proliferation, cell expansion, and cell death processes, TCTP was reported to interact with several proteins such as the 151 tumor suppressor p53, BAX (Bcl2 Associated X protein), or MDM2 (Mouse 152 Double Minute, 2000.
, embryo growth in the early stages of development and thus in agreement 210 with Brioudes et al. (2010) (Fig. 7.3b). Moreover, the authors demonstrated that like 211 animal TCTPs, plant TCTP, even though lacking signal secretion peptide, can be 212 secreted to plant apoplastic spaces via exosomes, agreement with Brioudes et al. (2010), very 206 recently Hafidh et, vol.213, 2004.
, In these plants, TCTP downregulation leads to 225 delayed plant development and smaller organs compared to wild type. Moreover, 226 flowers were smaller and root growth was reduced (Tao et al. 2015; Bruckner et al. of cell proliferation, Arabidopsis AtTCTP protein was shown to accumulate 231 in highly dividing cells, Pisum sativum, mRNA was 232 localized predominantly in dividing cells of root caps and in other rapidly growing 233 tissues as young leaves and stems, vol.224, 1997.
, TCTP protein accumulation was correlated with the accumulation 235 of other cell proliferation proteins in the skin of young potato tubers, an actively 236 dividing tissue, 2008.
, To the best of our knowledge, only two studies showed the direct implication of 238 TCTP in the control of cell proliferation. In a first study, Brioudes et al.(2010) used 239 synchronized tobacco (Nicotiana tabacum) BY-2 cells knockdown for NtTCTP to B
, TCTP prevents PCD is still unknown as counterparts of TCTP interacting mam
, As discussed above, TCTP is absolutely required for plant development and organ 288 size determination. In Arabidopsis and tobacco, the downregulation of TCTP leads 289 to severe developmental defects that are at least in part due to perturbation of cell 290 proliferation, but many other processes are affected. A number of published work 291 associates TCTP to other cellular functions and signaling molecules/pathways, 292 the next paragraph, we will provide the up-to-date information on the putative links 293 between TCTP and these cellular functions and signaling molecules/pathways
, Is TCTP a Component of the TOR Pathway?
, reported that Drosophila dTCTP controls organ growth by 296 positively regulating the TARGET OF RAPAMYCIN (TOR) pathway. TOR kinase 297 is part of a signaling complex that controls cell proliferation and growth in animals 298 and in plants, Deprost et al, 2004.
, In mammals, 301 the TOR pathway (mTOR) controls cell growth by acting as a central regulator of 302 protein synthesis and ribosome biogenesis at the transcriptional and translational 303 levels downstream mitogens and nutrients sensing and by integrating signaling 304 pathways ( AU4 Wullschleger et al. 2006). mTOR activity is positively controlled by 305 the small Ras GTPase, Rheb, that binds directly to mTOR kinase domain in order to 306 activate the mTOR complex in a GTP-dependent manner, The Tuberous Sclerosis Complex (TSC) negatively regulates TOR pathway 308 by inactivating, vol.307, 2000.
, In this model, dTCTP directly associates with dRheb and is 314 required for its activation in vivo, which in turn positively controls TOR activity; 315 thus, TCTP may have an opposite function to TSC. Mouse TCTP was also reported 316 to function as GEF upstream of S6K (Chen et al. 2007) and human TCTP was 317 demonstrated to activate the mTOR pathway in vivo (Dong et al. 2009). In plants, 318 TOR is highly conserved and its expression positively correlates with growth 319, Drosophila 310 melanogaster, dTCTP acts in parallel to TSC, but upstream of dRheb, 2006.
, 2001) and that it can interact with 367 many cellular proteins involved in cell growth or survival control, it was tempting 368 to propose animal TCTP as a protein chaperone
, Most of the published work simply provided hypothesis based on observa381 tions, and sometimes results are contradictory. 382 In plants, auxin is an essential hormone involved in a wide variety of functions 383 such as cell division, organogenesis, senescence, apical dominance, gravitropism, 384 root growth, Whether TCTP acts as a GEF or not on Rheb proteins remains to be clarified, 2008.
, In 390 cabbage, TCTP-RNAi plants grow slower and accumulate low auxin contents 391 compared to the wild-type (Cao et al. 2010). The authors suggested that such 392 decrease of endogenous auxin content could be responsible of the observed delayed 393 growth. However, this remains highly hypothetical as no direct proof has been 394 provided by the authors to support such hypothesis, RNAi plants were less sensitive to increasing concentrations of 386 exogenous auxin compared to the wild-type and proposed an implication of TCTP 387 in the auxin signaling pathway, 1987.
, ABA is a plant hormone known to be implicated in seed 398 germination and response to drought. It has been suggested in Arabidopsis that 399 AtTCTP interacts with ABA signaling pathway to control stomatal closure mech400 anism in response to drought, Several studies reported interaction of TCTP with the abscisic acid (ABA) 397 signaling pathway, 2012.
, 404 reported that overexpression of Arabidopsis AtTCTP confers drought tolerance by 405 rapid ABA-mediated stomatal closure. The authors also proposed that this rapid 406 ABA-mediated stomatal closure is mediated by the interaction of AtTCTP with B. Léo et al. 429 interacts with the ethylene receptor NTHK1, and such interaction is enhanced by 430 ethylene and protects NTHK1 from degradation by the 26S proteasome and pro431 motes seedling growth (Tao et al. 2015). Some studies also report a change in TCTP 432 expression upon Jasmonic acid (JA) treatment. JA is a lipid-derived plant hormone 433 that regulates a wide range of processes, such as biotic and abiotic plant stress 434 responses (insects, pathogens, wounding.. .) and developmental processes (root 435 growth, senescence, Devoto and Turner, 2003.
, In cotton, overexpression of TCTP leads to important 438 decrease of expression of genes in the JA pathway, Hevea brasiliensis, HbTCTP expression is strongly reduced in response to MeJA 437 treatment, 2013.
, MeJA in Oryza sativa, These results are contradictory, and further experi, 2015.
, To summarize, we have little knowledge about the link between TCTP and 444 phytohormone signaling, and clearly, more studies are needed to determine the 445 role of TCTP in phytohormone signaling during plant growth and development
, Conversely to animals, plants have sessile lifestyle and they have to cope with and 448 adapt to their environment
, While some reports show that the expression of Arabidopsis AtTCTP and 451 tomato SlTCTP mRNAs is highly stable in different growth conditions (Brioudes 452 et al. 2010; Coker and Davies 2003), several other studies reported variations of 453 TCTP mRNA expression and/or protein accumulation in response to environmental 454 stresses (Fig. 7.4). In plants, TCTP expression was reported to vary in response to 455 salt stress, high temperature, drought, and pathogen attacks (Fig. 7.4). In the 456 following chapter, The availability of those elements directly impact their development, morphology, 450 and size
, Kim et al. (2012) showed 465 that in Arabidopsis, plants overexpressing AtTCTP were more tolerant to drought 466 due to reduced water loss. As described above, AtTCTP interacts with microtubules 467 in guard cells, which leads to microtubule depolymerization and rapid 468 ABA-mediated stomatal closure and reduced water loss, 2007.
, HR is one of the best-characterized defense responses in plants that occurs 524 during incompatible host-pathogen interaction. It is characterized by rapid, local525 ized PCD to restrict the spread of pathogens and protect the plant. Because HR is 526 one of the major defense mechanisms against pathogen attacks (Morel and Dangl 527 1997), TCTP is assumed to play a role in plants defense. Wide-genome as well as 528 proteomics analysis show that both TCTP transcript and protein were differentially 529 expressed and accumulated in response to attacks by viruses, bacteria, and fungi. 530 Analysis of proteomic changes in Arabidopsis infected by Pseudomonas 531 syringae demonstrated that infection with an avirulent strain or induction of HR 532 by bacteria correlates with TCTP downregulation, and conversely, TCTP is an anti-apoptotic protein associated with plant 522 hypersensitive response (HR) during incompatible interaction with many patho523 gens, 2006.
, Successful infection by Agrobacterium tumefaciens that requires 535 suppression of plant defense mechanism, also correlates with increased TCTP 536 expression (Veena et al. 2003). In agreement with these reports, NbTCTP silencing 537 in tobacco accelerates HR following infection by various bacteria, while TCTP 538 overexpression diminished HR response to bacterial infection, 2008.
, NtTCTP seems to negatively regulate HR cell death by acting on the MAPK540 regulated cell death pathway. Moreover, expression of the BAX protein in NtTCTP 541 downregulated plants accelerated the HR phenotype. BAX is a mammalian 542 pro-apoptotic protein, which in mammals antagonizes TCTP effect
, In a more recent study, 547 it was demonstrated that TCTP is an important host factor for an efficient infection 548 by potyviruses in tomato and Nicotiana benthamiana plants. Plants silenced for 549 TCTP exhibited reduced accumulation of PepYMV early during infection 550 (Bruckner et al. 2016). Furthermore, TCTP was observed in both nuclei and 551 cytoplasm of non-infected cells, while only in the cytoplasm of infected cells, TCTP was identified among the most strongly induced genes after 546 potyvirus infection in tomato, 2009.
, However, the mechanism by which TCTP promote successful infection by viruses 554 is not yet clear. In animals, the highly structured TCTP mRNA was shown to 555 activate the dsRNA-dependent protein kinase PKR (Bommer et al. 2002), thus 556 suggesting antiviral responses. In plants, dsRNA binding is used by plant viruses to 557 suppress RNA silencing, 2006.
, such as cell proliferation and cell death control or 605 response to abiotic and biotic stresses. The precise molecular mechanisms associ606 ated with many of these processes in plants are still unknown. In animals, TCTP is 607 involved in diverse cancers and is thought to be one of the best therapeutic targets to 608 fight against the disease (Acunzo et al. 2014). In plants, tumorogenesis occurs but 609 rarely induces physiological disorder (Doonan and Sablowski, plants as in animals, TCTP is involved and likely plays major roles in many 604 cellular processes (Fig. 7.4), 2010.
, However, to the best of our knowledge, only one report addressed the TCTP 612 functional conservation between plants and animals at the molecular level
, Biologie et Amélioration des Plantes" 619 Department of the French "Institut National de la Recherche Agronomique
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