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Cited by 106Plant Molecular Biology ResearchPlant Reproductive BiologyPlant Gene Expression Analysis

Divide et impera: boundaries shape the plant body and initiate new meristems

Open access
Wang, Quan, Hasson, Alice, Rossmann, Susanne, Theres, Klaus·2016·New Phytologist
DOI 10.1111/nph.13641
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Abstract

485 I. 485 II. 486 III. 491 IV. 491 V. 495 495 References 495 SUMMARY: Boundaries, established and maintained in different regions of the plant body, have diverse functions in development. One role is to separate different cell groups, for example the differentiating cells of a leaf primordium from the pluripotent cells of the apical meristem. Boundary zones are also established during compound leaf development, to separate young leaflets from each other, and in many other positions of the plant body. Recent studies have demonstrated that different boundary zones share similar properties. They are characterized by a low rate of cell divisions and specific patterns of gene expression. In addition, the levels of the plant hormones auxin and brassinosteroids are down-regulated in boundary zones, resulting in a low differentiation level of boundary cells. This feature seems to be crucial for a second important role of boundary zones, the formation of new meristems. The primary shoot meristem, as well as secondary and ectopic shoot meristems, initiate from boundary cells that exhibit competence for meristem formation.

AI snapshot

Written from the paper’s full text.

TL;DR

This review argues that boundary zones are not merely physical separators but active developmental modules that shape plant architecture and serve as sites for new meristems. It synthesizes genetic and hormonal evidence from flowering plants, especially tomato and Arabidopsis, to show shared boundary properties and conserved regulators.

Key findings

  • Boundary zones show reduced cell division and limited expansion, often appearing as grooves; examples include fewer divisions at the tomato shoot apical meristem–leaf primordium boundary, down-regulated DNA synthesis and cell-cycle genes in Arabidopsis boundary cells, and smaller, less differentiated cells with lower Histone H4 mRNA at distal leaflet boundaries in tomato and Cardamine pratensis.
  • Conserved NAM/CUC transcription factors (e.g., CUC1/2/3, petunia NAM, snapdragon CUP, tomato Goblet) are expressed in boundaries; their loss causes organ fusions and simpler compound leaves. Other regulators include LOF1/2, JLO, LOB/BOP, LAS/Ls/MOC1, RAX1/Blind, and ROX/LAX1/BA1.
  • Hormonal control is central: brassinosteroids via BZR1 repress CUC and LOF1; LOB and brassinosteroids form a negative feedback loop; boundaries are regions of low auxin signaling due to PIN1 repolarization; cytokinin promotes meristem initiation. In maize, auxin biosynthesis/transport mutants (spi1, vt2, bif2) affect axillary meristem formation, while in Arabidopsis low auxin in the leaf axil helps maintain competence.
  • Boundaries can initiate meristems: the inter-cotyledon boundary gives rise to the shoot apical meristem, leaf axils produce axillary meristems, and distal leaflet boundaries produce ectopic meristems in tomato. Timing examples include Arabidopsis Col-0 axillary meristems first detected in P16 leaf axils and tomato cv Moneymaker axillary meristems detected in P7 axils; not all boundaries form meristems, such as the proximal leaflet boundary.

Methods

  • Narrative review with no new experimental data; it synthesizes published genetic, molecular, histological, and hormonal studies across flowering plants.
  • Focus is on flowering plants, with examples from tomato, Arabidopsis, Cardamine hirsuta, maize, rice, snapdragon, petunia, Medicago truncatula, pea, and others.
  • Main analysis is conceptual integration of boundary establishment, leaf shaping, and meristem initiation, including a gene/hormone interaction network and a table of genes involved in boundary and meristem formation.

Limitations

  • The authors state that space limitations prevented inclusion of some publications and that the review focuses only on flowering plants.
  • The authors identify open questions: molecular interactions among regulators, whether expression of one boundary gene is sufficient to initiate boundary formation, how hormone signaling and gene expression affect each other, the physical properties of boundary zones, and the triggers that ultimately initiate meristem formation.
  • Evident from the text: as a review, it provides no primary data or systematic quantitative synthesis; conclusions rely on cited studies and models, and some mechanisms, such as PIN1 re-localization, remain unresolved.

Model-written summary. Check it against the paper before relying on it.

Integrity signals

Registry

Crossref · 10.1111/nph.13641

Cites

101 of 101 references verified0 citation errors · 0 not checkable

Statistics

Nothing to recompute.

Claims the paper makes

51 of 224 cited sources checked · 30 supported · 2 flagged · accuracy not yet measured

  • Premisecitation

    Loss-of-function mutants of LAS show fusion of lateral branches to the main axis.

    “| LAS | Ls (tomato), MOC1 (rice | GRAS | Fusion of lateral branches to | SAM and lateral organs, | Schumacher et al. (1999); Li et al. |”
    • Also stated in: § V. Conclusions/future perspectives · plain
      “| | (Oryza sativa)) | | the main axis, reduced | leaflet and rachis, in | (2003); Greb et al. (2003); Busch |”
    Quote in the paperSubstantial claim3 of 3 sources checked
    • cites Schumacher et al. (1999) source

      The paper identifies the tomato ls alleles as complete loss-of-function mutants (Discussion; page not provided). But it describes their vegetative phenotype as an absence of side-shoots because axillary cells fail to retain meristematic character, not fusion of lateral branches to the main axis (Abstract; page not provided). (LLM, conf=0.88)

      Checked by a reasoning model — may be wrong.

      Experimental — accuracy not yet measured
    • cites Greb et al. (2003) source
    • cites Busch et al. (2011) source

      The Results (page unavailable) say that Ls mutations manifest only as a lack of axillary meristems. The same section attributes side-shoot fusions to bl mutants, not Ls mutants. Although the Discussion broadly mentions organ fusions among boundary-gene mutants, it does not identify fusion of lateral branches to the main axis as an Ls phenotype. (LLM, conf=0.86)

      Checked by a reasoning model — may be wrong.

      Experimental — accuracy not yet measured
  • Premisecitation

    Boundary zones between the meristem and leaf primordia show similar properties to boundaries between leaflets.

    “Recent experiments have shown that the boundary zones between the meristem and leaf primordia show similar properties to the boundaries between leaflets (Busch et al., 2011)”
    Quote in the paperSubstantial claim1 of 1 sources checked
    • cites Busch et al., 2011 source
  • Premisecitation

    Distal leaflet boundaries of tomato and Cardamine pratensis leaves harbor smaller cells compared with surrounding tissues.

    “The distal leaflet boundaries of tomato and Cardamine pratensis leaves were shown to harbor smaller cells in comparison to surrounding tissues (Rossmann et al., 2015).”
    Quote in the paperNot a substantial claim1 of 1 sources checked
    • cites Rossmann et al., 2015 source
  • Premisecitation

    In Arabidopsis, boundary cells between inflorescence and floral meristems down-regulate DNA synthesis and expression of cell-cycle-related genes.

    “In Arabidopsis, boundary cells between inflorescence and floral meristems down-regulate DNA synthesis and expression of cell-cycle-related genes (Breuil-Broyer et al., 2004).”
    Quote in the paperNot a substantial claim1 of 1 sources checked
    • cites Breuil-Broyer et al., 2004 source
  • Premisecitation

    In wild type, BZR1 proteins accumulate to high levels in the central meristem and organ primordia, limiting expression of a specific set of genes, including CUC genes, to the boundary.

    “In wild type, BZR1 proteins accumulate to high levels in the central meristem and organ primordia, limiting the expression of a specific set of genes, including CUC genes, to the boundary (Gendron et al., 2012).”
    Quote in the paperSubstantial claim1 of 1 sources checked
    • cites Gendron et al., 2012 source
  • May combine two claims. 63
  • Classified as probably not a substantial claim. 12

  • May combine two claims.

    Premisecitation

    Cells at distal leaflet boundaries of tomato and Cardamine pratensis are less differentiated, lack differentiation markers such as stomata or trichomes, and exhibit down-regulation of Histone H4 mRNA accumulation, indicating a low cell division rate.

    “At the same time, these cells are less differentiated, as indicated by the lack of differentiation markers, such as stomata or trichomes, and exhibit a down-regulation of Histone H4 mRNA accumulation, indicating a low cell division rate (Rossmann et al., 2015; Fig. 1d).”
    Quote in the paperNot a substantial claim1 of 1 sources checked
    • cites Rossmann et al., 2015 source

      Split into 2 atoms — 0 confirmed, 2 partially supported, 0 disputed, 0 misattributed, 0 unverifiable. Expand the claim to see per-atom verdicts.

      Experimental — accuracy not yet measured

  • May combine two claims.

    Premisecitation

    These MYB genes are specifically expressed in the leaf axil and plants homozygous for loss-of-function alleles show fusions of axillary shoots to their subtending leaves or to the main axis, indicating the gene product is required for axillary meristem formation and organ separation.

    “These genes are specifically expressed in the leaf axil and plants homozygous for loss-of-function alleles show fusions of axillary shoots to their subtending leaves or to the main axis (Greb et al., 2003;Busch et al., 2011), indicating that the respective gene product is required not only for axillary meristem formation, but also for organ separation.”
    Quote in the paperSubstantial claim2 of 2 sources checked
    • cites Greb et al., 2003 source

      Split into 2 atoms — 1 confirmed, 1 partially supported, 0 disputed, 0 misattributed, 0 unverifiable. Expand the claim to see per-atom verdicts.

      Experimental — accuracy not yet measured
    • cites Busch et al., 2011 source

      Split into 2 atoms — 1 confirmed, 1 partially supported, 0 disputed, 0 misattributed, 0 unverifiable. Expand the claim to see per-atom verdicts.

      Experimental — accuracy not yet measured

  • May combine two claims.

    Premisecitation

    Auxin response maxima instruct the position of leaf primordia at the shoot apex and the pattern of leaflets during complex leaf development.

    “Auxin response maxima instruct, for example, the position of leaf primordia at the shoot apex (Reinhardt et al., 2003;Bayer et al., 2009) and the pattern of leaflets during complex leaf development (Barkoulas et al., 2008;Koenig et al., 2009;Ben-Gera et al., 2012).”
    Quote in the paperSubstantial claim1 of 3 sources checked
    • cites Reinhardt et al., 2003 source
    • cites Bayer et al., 2009 source
    • cites Barkoulas et al., 2008 source

  • May combine two claims.

    Premisecitation

    abcb19 mutants displayed increased auxin concentrations in boundaries of the inflorescence meristem, whereas CUC2 and LOF1 expression was decreased.

    “abcb19 mutants displayed increased auxin concentrations in boundaries of the inflorescence meristem, whereas CUC2 and LOF1 expression was found to be decreased (Zhao et al., 2013).”
    Quote in the paperSubstantial claim1 of 1 sources checked
    • cites Zhao et al., 2013 source

  • May combine two claims.

    Premisecitation

    Auxin application leads to the formation of ectopic leaflets and lamina outgrowth, suggesting auxin triggers leaflet initiation and lamina growth.

    “In contrast, auxin application leads to the formation of ectopic leaflets and lamina outgrowth, suggesting that auxin triggers leaflet initiation and lamina growth (Barkoulas et al., 2008;Koenig et al., 2009;Ben-Gera et al., 2012).”
    Quote in the paperSubstantial claim1 of 3 sources checked
    • cites Barkoulas et al., 2008 source
    • cites Koenig et al., 2009 source
    • cites Ben-Gera et al., 2012 source

  • May combine two claims.

    Premisecitation

    Young e mutant leaf primordia show a broad auxin response at the leaf margin, corroborating that the interspersed pattern of auxin response maxima in wild-type tomato plants is a prerequisite for the observed pattern of compound leaf development.

    “Young e mutant leaf primordia show a broad auxin response at the leaf margin, corroborating that the interspersed pattern of auxin response maxima in wild-type tomato plants is a prerequisite for the observed pattern of compound leaf development (Koenig et al., 2009;Ben-Gera et al., 2012).”
    Quote in the paperSubstantial claim1 of 2 sources checked
    • cites Koenig et al., 2009 source
    • cites Ben-Gera et al., 2012 source

      Split into 2 atoms — 1 confirmed, 1 partially supported, 0 disputed, 0 misattributed, 0 unverifiable. Expand the claim to see per-atom verdicts.

      Experimental — accuracy not yet measured

  • May combine two claims.

    Interpretationcitation

    The leaf axil and the distal leaflet boundary harbor pluripotent cell groups that retain the competence for meristem formation.

    “The patterns of cell division and the absence of well-known differentiation markers, such as stomata and trichomes, indicate that the leaf axil and the distal leaflet boundary harbor pluripotent cell groups that retain the competence for meristem formation (Rossmann et al., 2015).”
    Quote in the paperSubstantial claim1 of 1 sources checked
    • cites Rossmann et al., 2015 source

  • May combine two claims.

    Premisecitation

    c mutants develop less complex leaves than wild-type, with a reduced number of leaflets and a smooth leaflet margin.

    “In comparison to wild-type, c mutants develop less complex leaves (Busch et al., 2011), with a reduced number of leaflets and a smooth leaflet margin.”
    Quote in the paperSubstantial claim1 of 1 sources checked
    • cites Busch et al., 2011 source

  • May combine two claims.

    Premisecitation

    Potato leaf is expressed in the boundaries between initiating leaflets and represses growth in these regions.

    “Potato leaf is expressed in the boundaries between initiating leaflets and represses growth in these regions (Busch et al., 2011).”
    Quote in the paperSubstantial claim1 of 1 sources checked
    • cites Busch et al., 2011 source

      The Results section reports that Potato leaf (C) is expressed at the proximal and distal boundaries of young leaflet primordia. The Discussion proposes that C may repress growth in boundary regions, based partly on growth retardation when C is misexpressed. The claim presents this proposed function as established. (LLM, conf=0.88)

      Experimental — accuracy not yet measured

  • May combine two claims.

    Premisecitation

    LAS is expressed in the boundary between the SAM and lateral organs, leaflet and rachis, and in mature tiller bud.

    “| LAS | Ls (tomato), MOC1 (rice | GRAS | Fusion of lateral branches to | SAM and lateral organs, | Schumacher et al. (1999); Li et al. |”
    • Also stated in: § V. Conclusions/future perspectives · plain
      “| | (Oryza sativa)) | | the main axis, reduced | leaflet and rachis, in | (2003); Greb et al. (2003); Busch |”
    Quote in the paperSubstantial claim3 of 3 sources checked
    • cites Schumacher et al. (1999) source

      The Results section (‘RT-PCR Detection of Ls mRNA’; page unavailable) reports Ls transcripts in tomato shoot tips and young leaves. The Discussion says the authors could not analyze expression at the in situ level. The paper therefore does not establish expression specifically at the SAM–lateral-organ boundary, between leaflet and rachis, or in mature tiller buds. (LLM, conf=0.83)

      Experimental — accuracy not yet measured
    • cites Greb et al. (2003) source

      The section “LAS transcripts accumulate in the axils of all primordia derived from the SAM” reports LAS expression at the boundary between Arabidopsis leaf primordia and the shoot apex. The paper does not report LAS expression between leaflet and rachis or in mature tiller buds. (LLM, conf=0.88)

      Experimental — accuracy not yet measured
    • cites Busch et al. (2011) source

      The Introduction (page not available) places tomato Ls expression at the boundary between the leaf primordium and stem. The Results (page not available) report Ls mRNA in the leaf axil and at the distal boundary of leaflets, supporting the boundary-expression portion of the claim. The paper does not report LAS or its rice counterpart MOC1 being expressed in mature tiller buds. (LLM, conf=0.88)

      Experimental — accuracy not yet measured

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    The Arabidopsis LAS gene encodes a GRAS protein family protein.

    “| LAS | Ls (tomato), MOC1 (rice | GRAS | Fusion of lateral branches to | SAM and lateral organs, | Schumacher et al. (1999); Li et al. |”
    Quote in the paperNot a substantial claim2 of 3 sources checked
    • cites Schumacher et al. (1999) source

      The Results ('Ls Is a New Member of the VHIID Family'; page not provided) identify tomato Ls as a member of the VHIID protein family and mention a highly similar, unpublished Arabidopsis clone. The copy does not identify that clone as Arabidopsis LAS or establish that the Arabidopsis LAS gene encodes a GRAS-family protein. (LLM, conf=0.78)

      Experimental — accuracy not yet measured
    • cites Li et al. (2003)

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

    • cites Greb et al. (2003) source

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    The Arabidopsis RAX1 gene encodes a MYB protein family protein.

    “| RAX1 | Bl (tomato) | MYB | Reduced number of AMs | SAM and leaf primordia | Keller et al. (2006); M€ uller et al. |”
    Quote in the paperNot a substantial claim1 of 2 sources checked
    • cites Keller et al. (2006) source
    • cites Müller et al. (2006) source

  • May combine two claims.

    Interpretationreasoning

    Boundary zones are more than a physical separation layer between neighboring parts of the plant body; they provide a versatile module utilized in different developmental processes.

    “These observations already indicate that boundary zones are more than a physical separation layer between neighboring parts of the plant body; rather they provide a versatile module that is utilized in different developmental processes.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Premisecitation

    Boundaries are generally characterized by a low number of cell divisions and specific patterns of gene expression.

    “In general, boundaries are characterized by a low number of cell divisions (Hussey, 1971;Reddy et al., 2004) and specific patterns of gene expression (Souer et al., 1996;Aida et al., 1997;Takada et al., 2001;Vroemen et al., 2003;Weir et al., 2004;Berger et al., 2009).”
    Quote in the paperNot a substantial claim0 of 3 sources checked
    • cites Hussey, 1971 source
    • cites Reddy et al., 2004 source
    • cites Souer et al., 1996 source

  • May combine two claims.

    Premisecitation

    In Arabidopsis, small cells accumulate at the sinuses of leaf teeth, whereas adjacent cells along a tooth adopt an elongated shape.

    “In Arabidopsis, small cells accumulate at the sinuses of leaf teeth, whereas adjacent cells (i.e.along a tooth) adopt an elongated shape (Kawamura et al., 2010).”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites Kawamura et al., 2010 source

  • May combine two claims.

    Premisecitation

    NAM/CUC family genes are consistently expressed in boundary zones and their loss of function causes fusion of adjacent organs that would normally be separated.

    “These genes are consistently expressed in boundary zones and their loss of function causes fusion of adjacent organs (e.g.cotyledons, floral organs, and ovules) that would normally be separated (Souer et al., 1996;Aida et al., 1997;Takada et al., 2001;Vroemen et al., 2003;Weir et al., 2004; Fig. 1e,f).”
    Quote in the paperSubstantial claim0 of 3 sources checked
    • cites Souer et al., 1996 source
    • cites Aida et al., 1997 source
    • cites Takada et al., 2001 source

  • May combine two claims.

    Premisecitation

    jlo loss-of-function mutants show fusions of cotyledons and of floral organs.

    “jlo loss-offunction mutants show fusions of cotyledons and of floral organs.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Premisenone given

    High BR concentrations promote cell enlargement, whereas low BR concentrations cause a reduction in cell size.

    “High BR concentrations promote cell enlargement, whereas low BR concentrations cause a reduction in cell size.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Premisecitation

    The NAM/CUC ortholog from Medicago truncatula is necessary for organ separation including compound leaf development, and is also required for floral organ identity and development.

    “Recently, it has been reported that the NAM/CUC ortholog from Medicago truncatula is necessary for organ separation including compound leaf development, and is also required for floral organ identity and development (Cheng et al., 2012).”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites Cheng et al., 2012 source

  • Classified as probably not a substantial claim.

    Premisecitation

    NAM/CUC proteins exert their function by repressing growth in boundary regions.

    “It has been proposed that NAM/CUC proteins exert their function by repressing growth in the boundary regions, because mutations in these genes or down-regulation of their expression leads to ectopic growth in the boundaries between adjacent morphological structures (Souer et al., 1996”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Souer et al., 1996 source

  • May combine two claims.

    Premisecitation

    bop1 bop2 double mutants show ectopic tissue growth from the lamina and lack a distinct petiole.

    “bop1 bop2 double mutants show ectopic tissue growth from the lamina, and lack a distinct petiole (Norberg et al., 2005;Ha et al., 2007).”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Norberg et al., 2005 source
    • cites Ha et al., 2007

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • May combine two claims.

    Premisecitation

    Both bop1 and bop2 mutants display defects in inflorescence development, frequently forming fused inflorescences and generating multiple flowers from the same node.

    “Both bop1 and bop2 mutants display defects in inflorescence development, as they frequently form fused inflorescences and generate multiple flowers from the same node (Hepworth et al., 2005;Ha et al., 2007).”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Hepworth et al., 2005 source
    • cites Ha et al., 2007

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • May combine two claims.

    Interpretationreasoning

    The functions of these boundary genes are well conserved through evolution and have been used in a modular way for the development of various structures in the plant body.

    “Their functions are well conserved through evolution and have been used in a modular way for the development of various structures in the plant body.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Premisereasoning

    Ectopic expression of LOB causes a reduced BR response, and expression of BAS1 under the LOB promoter can rescue the organ fusion defects of the lob mutant.

    “Ectopic expression of LOB causes a reduced BR response, and expression of BAS1 under the control of the LOB promoter can rescue the organ fusion defects of the lob mutant.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Premisecitation

    There is no evidence that CUC gene expression is altered in the bop1 bop2 double mutant, suggesting BOP1/2 are not playing a role in the initial establishment of boundary domains.

    “There is no evidence that CUC gene expression is altered in the bop1 bop2 double mutant, suggesting that BOP1/2 are not playing a role in the initial establishment of boundary domains (Khan et al., 2014).”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites Khan et al., 2014 source

  • May combine two claims.

    Premisecitation

    LOB is activated by BOP1 and BOP2 genes, which are transiently expressed in leaf primordia and restricted to the boundary when primordia first appear morphologically distinct from the meristem.

    “LOB is activated by BLADE-ON-PETIOLE1 (BOP1) and BOP2 genes, which are transiently expressed in leaf primordia and restricted to the boundary when primordia first appear as morphologically distinct from the meristem (Ha et al., 2004(Ha et al., , 2007;;Norberg et al., 2005).”
    Quote in the paperNot a substantial claim0 of 3 sources checked
    • cites Ha et al., 2004

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

    • cites Ha et al., 2007

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

    • cites Norberg et al., 2005 source

  • May combine two claims.

    Premisecitation

    As a consequence, CUC gene expression is not restricted to the boundary region between the two cotyledons, but expands throughout the top of the embryo and represses cotyledon formation.

    “As a consequence, CUC gene expression is not restricted to the boundary region between the two cotyledons, but expands throughout the top of the embryo and represses cotyledon formation.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Premisecitation

    In the C. hirsuta pin1 mutant, both leaf primordium initiation at the shoot apex and lateral leaflet formation are disrupted.

    “In the C. hirsuta pin1 mutant, both leaf primordium initiation at the shoot apex and lateral leaflet formation are disrupted (Barkoulas et al., 2008).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Barkoulas et al., 2008 source

  • May combine two claims.

    Premisecitation

    Auxin application to the cuc1 single mutant significantly enhanced the cotyledon fusion phenotype, corroborating that auxin represses CUC2 expression.

    “In addition, auxin application to the cuc1 single mutant significantly enhanced the cotyledon fusion phenotype, corroborating that auxin represses CUC2 expression.”
    Quote in the paperSubstantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

    Premisecitation

    ARF activity is down-regulated by interaction with AUX/IAA proteins, which are targeted for degradation by the proteasome in the presence of auxin.

    “ARF activity is down-regulated by interaction with AUXIN/INDOLE ACETIC ACID (AUX/IAA) proteins (Liscum & Reed, 2002;Leyser, 2006), which are targeted for degradation by the proteasome in the presence of auxin (Dharmasiri et al., 2005;Kepinski & Leyser, 2005).”
    Quote in the paperNot a substantial claim0 of 3 sources checked
    • cites Liscum & Reed, 2002 source
    • cites Leyser, 2006 source
    • cites Dharmasiri et al., 2005 source

  • May combine two claims.

    Premisecitation

    CUC2 promotes the formation of auxin maxima through its action on PIN1 re-orientation and in return auxin downregulates CUC2 expression, forming a robust feedback loop.

    “CUC2 promotes the formation of auxin maxima through its action on PIN1 re-orientation and in return auxin downregulates CUC2 expression, forming a robust feedback loop (Bilsborough et al., 2011).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Bilsborough et al., 2011 source

  • May combine two claims.

    Premisecitation

    The shift in auxin transport direction leads to auxin depletion and enables boundary zone formation between the leaf primordium and the meristem.

    “This shift in auxin transport direction leads to auxin depletion and enables boundary zone formation between the leaf primordium and the meristem (Wang et al., 2014a,b).”
    Quote in the paperSubstantial claim0 of 2 sources checked
    • cites Wang et al., 2014a

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

    • cites Wang et al., 2014b

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • May combine two claims.

    Premisecitation

    In pid mutants, normal PIN1 polarization is disrupted, resulting in abnormal auxin distribution.

    “In pid mutants, normal PIN1 polarization is disrupted, resulting in abnormal auxin distribution (Wang et al., 2014a).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Wang et al., 2014a

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • May combine two claims.

    Premisecitation

    In several compound-leaf species, CUC/GOB orthologous genes are expressed in the boundary of leaflets and down-regulation of their expression results in simplification of leaf architecture.

    “In several compound-leaf species, CUC/GOB orthologous genes are expressed in the boundary of leaflets and down-regulation of their expression results in simplification of leaf architecture (Blein et al., 2008;Berger et al., 2009;Cheng et al., 2012; Fig. 2d).”
    Quote in the paperSubstantial claim0 of 3 sources checked
    • cites Blein et al., 2008 source
    • cites Berger et al., 2009 source
    • cites Cheng et al., 2012 source

  • May combine two claims.

    Premisereasoning

    CUC2 expression conditions a repression of growth, whereas elevated auxin concentrations promote outgrowth of the leaf margin.

    “CUC2 expression conditions a repression of growth, whereas elevated auxin concentrations promote outgrowth of the leaf margin.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Premisecitation

    This patterning is dependent on PIN1 and CUC2 activity, as pin1 and cuc2 mutants develop smooth leaf margins.

    “This patterning is dependent on PIN1 and CUC2 activity, as pin1 and cuc2 mutants develop smooth leaf margins.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Premisecitation

    Boundaries between cotyledons and between developing leaflets are correlated with low auxin concentrations.

    “Furthermore, the boundaries between cotyledons (Furutani et al., 2004) and between developing leaflets (Koenig et al., 2009) have been correlated with low auxin concentrations.”
    Quote in the paperSubstantial claim0 of 2 sources checked
    • cites Furutani et al., 2004 source
    • cites Koenig et al., 2009 source

  • May combine two claims.

    Interpretationcitation

    Re-orientation of PIN1 and re-orientation of microtubule arrays do not directly depend on each other, but seem to be regulated both by an unknown upstream mechanism.

    “However, re-orientation of PIN1 and re-orientation of microtubule arrays do not directly depend on each other, but seem to be regulated both by an unknown upstream mechanism.”
    Quote in the paperNot a substantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    Auxin regulates transcript accumulation of target genes by activation of ARFs.

    “Auxin regulates transcript accumulation of target genes by activation of AUXIN RESPONSE FACTORS (ARFs) (Liscum & Reed, 2002;Leyser, 2006).”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Liscum & Reed, 2002 source
    • cites Leyser, 2006 source

  • May combine two claims.

    Premisereasoning

    The developmental timing of meristem initiation depends on the type of meristem, species-specific or even accession-specific patterns of meristem formation, and environmental conditions.

    “The developmental timing of meristem initiation depends on the type of meristem, species-specific or even accession-specific patterns of meristem formation, and environmental conditions.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Premisecitation

    CUC/GOB genes restrict growth in the boundary zones between initiating leaflets and promote the development of separated units in a complex leaf.

    “Similar to their activity at the margin of Arabidopsis leaves, CUC/GOB genes restrict growth in the boundary zones between initiating leaflets and promote, thereby, the development of separated units in a complex leaf.”
    Quote in the paperSubstantial claim0 of 3 sources checked
    • cites Blein et al., 2008 source
    • cites Berger et al., 2009 source
    • cites Cheng et al., 2012 source

  • May combine two claims.

    Premisenone given

    EMs initiate from the distal, but not from the proximal leaflet boundary.

    “EMs initiate from the distal, but not from the proximal leaflet boundary (Fig. 4c,f).”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Premisecitation

    RAX1 is specifically expressed in the boundary region between SAM and leaf primordia and modulates AM formation during early vegetative development in Arabidopsis.

    “Another R2R3 MYB protein, called REGULATOR OF AXILLARY MERISTEMS1 (RAX1), which is also specifically expressed in the boundary region between SAM and leaf primordia, modulates AM formation during early vegetative development in Arabidopsis (Keller et al., 2006;M€ uller et al., 2006), whereas the orthologous genes in pepper (Capsicum annuum) and tomato regulate both AM initiation and transition to”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Keller et al., 2006 source
    • cites M€ uller et al., 2006

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • May combine two claims.

    Premisecitation

    LOF1 and its paralog LOF2 are both specifically expressed in the adaxial leaf boundary and modulate AM formation in Arabidopsis.

    “Furthermore, LOF1 and its paralog LOF2 are both specifically expressed in the adaxial leaf boundary and modulate AM formation in Arabidopsis (Lee et al., 2009).”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites Lee et al., 2009 source

  • May combine two claims.

    Premisecitation

    RCO transcripts accumulate at the base of terminal and lateral leaflets, but are absent from the meristem-leaf boundary, suggesting that it does not represent a typical boundary-specific gene.

    “RCO transcripts accumulate at the base of terminal and lateral leaflets, but are absent from the meristem-leaf boundary, suggesting that it does not represent a typical boundary-specific gene (Vlad et al., 2014).”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites Vlad et al., 2014 source

  • May combine two claims.

    Premisenone given

    Several key regulators of boundary formation share a boundary-specific expression pattern and conspicuous organ fusions in their loss-of-function mutants.

    “Several key regulators of boundary formation have been identified that share the features of a boundary-specific expression pattern and conspicuous organ fusions in their loss-of-function mutants (Fig. 3; Table 1).”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Premisecitation

    The primary SAM initiates at the bending-cotyledon stage from a small group of cells and bulges into the characteristic dome-shaped structure.

    “The primary SAM initiates at the bending-cotyledon stage from a small group of cells (Barton & Poethig, 1993) and bulges into the characteristic domeshaped structure (Fig. 4a,d).”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites Barton & Poethig, 1993 source

  • May combine two claims.

    Premisecitation

    After germination, leaf primordia develop from the peripheral zone of the SAM, establishing new boundary zones between leaf primordia and the SAM.

    “After germination, leaf primordia develop from the peripheral zone of the SAM (Barton, 2010), establishing new boundary zones between leaf primordia and the SAM.”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Barton, 2010 source

  • May combine two claims.

    Interpretationcitation

    A low auxin environment is required for AM formation in Arabidopsis, whereas in maize auxin maxima are associated with the formation of AMs.

    “These results suggest that a low auxin environment is required for AM formation in Arabidopsis, whereas in maize auxin maxima are associated with the formation of AMs.”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Wang et al., 2014a

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • May combine two claims.

    Premisecitation

    LAX1 and BA1 are key regulators of shoot and inflorescence branching in rice and maize, respectively.

    “LAX PANICLE1 (LAX1) and BARREN STALK1 (BA1) were characterized as key regulators of shoot and inflorescence branching in the monocots rice and maize, respectively (Komatsu et al., 2003;Gallavotti et al., 2004).”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Komatsu et al., 2003 source
    • cites Gallavotti et al., 2004 source

  • May combine two claims.

    Premisecitation

    EM formation correlates with and is dependent on specific expression of the transcriptional regulators Ls and Gob.

    “This EM formation correlates with and is dependent on specific expression of the transcriptional regulators Ls and Gob.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Premisecitation

    CUC1 and CUC2 are required for the formation of the inter-cotyledon boundary zone and the SAM.

    “Very important are the NAM/CUC genes CUC1 and CUC2, required for the formation of the inter-cotyledon boundary zone and the SAM.”
    Quote in the paperSubstantial claim0 of 2 sources checked
    • cites Aida et al., 1997 source
    • cites Aida et al., 1999 source

  • May combine two claims.

    Interpretationcitation

    High CK concentrations promote leaflet formation, while low CK concentrations restrict it.

    “These data suggest that high CK concentrations promote leaflet formation, while low CK concentrations restrict it.”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Shani et al., 2010 source

  • May combine two claims.

    Interpretationcitation

    Mutations in either one of the auxin biosynthesis genes spi1 or vt2 strongly compromise AM formation, underpinning a crucial role for local auxin biosynthesis in this process.

    “Mutations in either one of these auxin biosynthesis genes strongly compromise AM formation, underpinning a crucial role for local auxin biosynthesis in this process.”
    Quote in the paperSubstantial claim0 of 2 sources checked
    • cites Gallavotti et al., 2008a

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

    • cites Phillips et al., 2011 source

  • May combine two claims.

    Premisecitation

    In Arabidopsis leaf axils, a CK pulse precedes AM formation and CK perception and signaling mutants are compromised in AM development.

    “Recently, it was shown that in Arabidopsis leaf axils a CK pulse precedes AM formation and that CK perception and signaling mutants are compromised in AM development (Wang et al., 2014b).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Wang et al., 2014b

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • May combine two claims.

    Interpretationreasoning

    Similar gene modules, involving two paralogous MYB genes (Bl and C), Gob and Ls, are utilized in both boundaries and seem to be required for full functionality of these zones.

    “Similar gene modules, involving two paralogous MYB genes (Bl and C), Gob and Ls, are utilized in both boundaries and seem to be required for full functionality of these zones.”
    Quote in the paperSubstantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    The Arabidopsis JLO gene encodes an LBD protein family protein.

    “| JLO | Unknown | LBD | Fusion of cotyledons and | SAM and organ primordia | Borghi et al. (2007); Bureau et al. |”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Borghi et al. (2007) source
    • cites Bureau et al. (2010) source

  • May combine two claims.

    Premisecitation

    Gob in tomato, CUP in snapdragon, and NAM in petunia are homologous to Arabidopsis CUC1-3.

    “| CUC1-3 | Gob (tomato (Solanum | NAC | Fusions of various organs, | SAM and lateral organs | Souer et al. (1996); Aida et al. |”
    • Also stated in: § V. Conclusions/future perspectives · plain
      “| | (Petunia hybrida)) | | AMs | leaves | et al. (2009) |”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Souer et al. (1996) source
    • cites Weir et al. (2004) source

  • May combine two claims.

    Premisecitation

    CK biosynthesis is required for axillary bud formation, as AM initiation is strongly reduced in the early vegetative phase of ipt3 ipt5 ipt7 triple mutants.

    “It was also shown that CK biosynthesis is required for axillary bud formation, as AM initiation is strongly reduced in the early vegetative phase of ipt3 ipt5 ipt7 triple mutants (M€ uller et al., 2015).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites M€ uller et al., 2015

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • May combine two claims.

    Premisecitation

    CUC1-3 is expressed in the boundary between the SAM and lateral organs (cotyledons, leaves, floral organs), leaflet and rachis, and emerging teeth of simple leaves.

    “| CUC1-3 | Gob (tomato (Solanum | NAC | Fusions of various organs, | SAM and lateral organs | Souer et al. (1996); Aida et al. |”
    • Also stated in: § V. Conclusions/future perspectives · plain
      “| | (snapdragon (Antirrhinum | | leaf species, smoother leaf | organs), leaflet and rachis, | Vroemen et al. (2003); Weir et al. |”
    Quote in the paperSubstantial claim0 of 3 sources checked
    • cites Souer et al. (1996) source
    • cites Vroemen et al. (2003) source
    • cites Berger et al. (2009) source

  • May combine two claims.

    Premisecitation

    CKs promote cell division in the shoot and positively regulate SAM size and activity.

    “CKs are known to promote cell division in the shoot and to positively regulate SAM size and activity (Barton, 2010).”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites Barton, 2010 source

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    The Arabidopsis BOP1 and BOP2 genes encode BTB/POZ protein family proteins.

    “| BOP1-2 | NOOT (medicago (Medicago | BTB/ | Fused inflorescences, simpler | SAM and leaf primordia, | Ha et al. (2004, 2007); Norberg et al. |”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Ha et al. (2004, 2007)

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

    • cites Norberg et al. (2005) source

  • May combine two claims.

    Premisecitation

    BOP1-2 is expressed in the boundary between the SAM and leaf primordia, and within the leaf.

    “| BOP1-2 | NOOT (medicago (Medicago | BTB/ | Fused inflorescences, simpler | SAM and leaf primordia, | Ha et al. (2004, 2007); Norberg et al. |”
    • Also stated in: § V. Conclusions/future perspectives · plain
      “| | truncatula)), COCH (pea | POZ | leaves in compound leaf | within the leaf | (2005); Couzigou et al. (2012); |”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Ha et al. (2004, 2007)

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

    • cites Norberg et al. (2005) source

  • May combine two claims.

    Interpretationreasoning

    The axillary boundary is characterized by low auxin and brassinosteroid signaling, and this may also hold for other boundary zones.

    “In addition, the axillary boundary is characterized by low auxin and brassinosteroid signaling, and, taking into account the other observed similarities, this may also hold for other boundary zones.”
    Quote in the paperNot a substantial claim

  • May combine two claims.

    Interpretationcitation

    The primary SAM and secondary shoot meristems originate from boundaries, and this connection is more widespread than anticipated.

    “Although it had already been known for a while that the primary SAM and secondary shoot meristems originate from boundaries, a new study has demonstrated that this connection is more widespread than anticipated.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Premisecitation

    NOOT in Medicago truncatula, COCH in pea, and Cul4 in barley are homologous to Arabidopsis BOP1-2.

    “| BOP1-2 | NOOT (medicago (Medicago | BTB/ | Fused inflorescences, simpler | SAM and leaf primordia, | Ha et al. (2004, 2007); Norberg et al. |”
    • Also stated in: § V. Conclusions/future perspectives · plain
      “| | (Pisum sativum)), Cul4 | | species | | Tavakol et al. (2015) |”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Couzigou et al. (2012) source
    • cites Tavakol et al. (2015) source

  • May combine two claims.

    Premisecitation

    LOF1-2 is expressed in the boundary between the stem and cauline leaves, inflorescence meristem and young floral primordia, and floral organs.

    “| LOF1-2 | T f(tomato) | MYB | Organ fusions, reduced | Stem and cauline leaves, | Lee et al. (2009); Naz et al. (2013) |”
    • Also stated in: § V. Conclusions/future perspectives · plain
      “| | | | species | primordia, floral organs | |”
    Quote in the paperSubstantial claim0 of 2 sources checked
    • cites Lee et al. (2009) source
    • cites Naz et al. (2013) source

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    The Arabidopsis LOF1 and LOF2 genes encode MYB protein family proteins.

    “| LOF1-2 | T f(tomato) | MYB | Organ fusions, reduced | Stem and cauline leaves, | Lee et al. (2009); Naz et al. (2013) |”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Lee et al. (2009) source
    • cites Naz et al. (2013) source

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    The Arabidopsis ROX gene encodes a bHLH protein family protein.

    “| ROX | ba1 (maize) | bHLH | Reduced number of AMs, | SAM and leaf primordia | Komatsu et al. (2003); Gallavotti |”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Komatsu et al. (2003) source
    • cites Gallavotti et al. (2004) source

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    The Arabidopsis LOB gene encodes an LBD protein family protein.

    “| LOB | ra2 (maize (Zea mays)) | LBD | Fusion of axillary branch and | SAM and lateral organs | Shuai et al. (2002); Zhou et al. |”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Shuai et al. (2002) source
    • cites Bortiri et al. (2006) source

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    The Arabidopsis LSH3 and LSH4 (OBO1/4) genes encode ALOG protein family proteins.

    “| LSH3/4 | G1 (rice) | ALOG | Only gain-of-function | SAM and lateral organs | Takeda et al. (2011); Cho & Zam- |”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Takeda et al. (2011) source
    • cites Cho & Zambryski (2011) source

  • May combine two claims.

    Premisecitation

    ra2 in maize, ELP/PLP in medicago, SLP in lotus, and APU in pea are homologous to Arabidopsis LOB.

    “| LOB | ra2 (maize (Zea mays)) | LBD | Fusion of axillary branch and | SAM and lateral organs | Shuai et al. (2002); Zhou et al. |”
    • Also stated in: § V. Conclusions/future perspectives · plain
      “| | SLP (lotus (Lotus japonicus)) | | | | et al. (2012) |”
    Quote in the paperSubstantial claim0 of 2 sources checked
    • cites Bortiri et al. (2006) source
    • cites Chen et al. (2012) source

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    Only a gain-of-function phenotype has been described for LSH3/4.

    “| LSH3/4 | G1 (rice) | ALOG | Only gain-of-function | SAM and lateral organs | Takeda et al. (2011); Cho & Zam- |”
    • Also stated in: § V. Conclusions/future perspectives · plain
      “| (OBO1/ | | | phenotype described | (cotyledons, leaves, floral | bryski (2011); Yoshida et al. (2009) |”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Takeda et al. (2011) source
    • cites Cho & Zambryski (2011) source

  • May combine two claims.

    Premisecitation

    ba1 in maize and LAX1 in rice are homologous to Arabidopsis ROX.

    “| ROX | ba1 (maize) | bHLH | Reduced number of AMs, | SAM and leaf primordia | Komatsu et al. (2003); Gallavotti |”
    • Also stated in: § V. Conclusions/future perspectives · plain
      “| | LAX1 (rice) | | reduced number of tillers | | et al. (2004); Yang et al. (2012) |”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Gallavotti et al. (2004) source
    • cites Yang et al. (2012) source
74 of 74

  • The paper relies on this without support.

    Little is known about the different roles of boundary zones in the complex process of shaping plant architecture.

    “To date, little is known about the different roles of boundary zones in the complex process of shaping plant architecture.”

  • The paper relies on this without support.

    The molecular mechanism by which NAM/CUC regulators control cell proliferation and differentiation in boundaries is not understood.

    “However, the molecular mechanism by which these regulators control cell proliferation and differentiation in the boundaries is still not understood.”

  • The paper relies on this without support.

    A detailed mechanistic explanation for intracellular re-shuffling of PIN proteins is missing.

    “A detailed mechanistic explanation for intracellular re-shuffling of PIN proteins is still missing.”
3 of 3

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Authors
Wang, Q., Hasson, A., Rossmann, S., & Theres, K.
Affiliations
Max Planck Institute for Plant Breeding Research (DE)
Corresponding author
Theres, Klaus
ORCID
Venue
New Phytologist
Publisher
Wiley
Published
2015-09-22
Volume
209
Issue
2
Pages
485-498
ISSN
  • 0028-646X (ISSN-L)
  • 1469-8137
DOI
10.1111/nph.13641
OpenAlex
W2100296318
Semantic Scholar
c2faf09b9cb227c5572adf205911fad988a97854
Cited by
106
References
98
Citations by year
2026: 6 · 2025: 7 · 2024: 9 · 2023: 4 · 2022: 10 · 2021: 16 · 2020: 18 · 2019: 10 · 2018: 7 · 2017: 10 · 2016: 7 · 2015: 1
FWCI
11.29
Type
journal article
Language
en
Open access
Licence
Locations
  • New Phytologist (publishedVersion)
  • PubMed (publishedVersion)
  • MPG.PuRe (Max Planck Society) (submittedVersion, other-oa)
  • Max Planck Digital Library (submittedVersion, other-oa)
  • MPG.PuRe (Max Planck Society) (submittedVersion)
Funders
  • Deutsche Forschungsgemeinschaft
  • Bundesministerium für Ernährung und Landwirtschaft
  • Max-Planck-Gesellschaft
  • Cluster of Excellence in Plant Sciences (1028)
Topics
  • Life Sciences › Agricultural and Biological Sciences › Plant Science › Plant Molecular Biology Research
  • Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Plant Reproductive Biology
  • Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Plant Gene Expression Analysis
Keywords
boundary region, plant development, shoot apical meristem, compound leaf development, leaf primordia, auxin, brassinosteroids, cell division rate
MeSH terms
Cell Division, Indoleacetic Acids, Gene Expression Regulation, Developmental, Plant Leaves, Meristem

About these checks

Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.

Claims read by deepseek-v4.1-flash under definition version 3 (extract-v3) on 2026-10-10.

Classified by jev-1.13-20260917 (gold-guideline-v0.2) on 2026-10-10.

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