Table 1.
Lesion mimic mutants.
| Mutant name | Gene ID | Gene product | PCD-related phenotypes | Possible mechanisms involved | References |
|---|---|---|---|---|---|
| acd1 | At3g44880 | Pheide a oxygenase involved in chlorophyll catabolic process | Lesions on older leaves, or when plants are submitted to a dark-light transfer | Accumulation of photoreactive pheide a | Greenberg and Ausubel, 1993; Gray et al., 1997; Tanaka et al., 2003 |
| lls1 (Zm) | |||||
| acd2 | At4g37000 | Red chlrorophyll catabolite reductase involved in chlorophyll catabolic process | Lesions on older leaves | Accumulation of red chlorophyll catabolites that causes singlet oxygen release | Mach et al., 2001; Yao and Greenberg, 2006; Pruzinska et al., 2007 |
| acd5 | At5g51290 | Ceramide kinase | Small restricted spontaneous lesions on leaves 5 weeks after planting | Accumulation of non-phosphorylated C2 ceramide | Greenberg et al., 2000; Liang et al., 2003; Bi et al., 2011 |
| acd6 (gof) | At4g14400 | Transmembrane protein with ankyrin domain | Yellowing leaves with cell death patches 2 weeks after planting | Constitutive activation of defense responses | Rate et al., 1999; Lu et al., 2003 |
| acd11 | At2g34690 | Protein that transfers Ceramide-1-Phosphate between membrane | Chlorosis that can engulfed the rosette | Ceramide and phytoceramide accumulation | Brodersen et al., 2002; Petersen et al., 2008; Simanshu et al., 2014 |
| agd2 | At4g33680 | Chloroplastic aminotransferase | Spotted necrotic lesions | Unknown | Rate and Greenberg, 2001; Song et al., 2004 |
| bir1 | At5g48380 | Receptor-like kinase belonging to the LRRX group | Lesions in cotyledons and true leaves | Activation of multiple defense signaling pathways | Gao et al., 2009 |
| cad1 | At1g29690 | Protein containing a MACPF domain | HR-like lesions on leaves | Unknown | Morita-Yamamuro et al., 2005 |
| camta3/sr1 | At2g22300 | Calmodulin-binding transcription factor | Chlorotic lesions on rosette leaves | Overexpression of immune regulator such as EDS1 and EIN3 | Yang and Poovaiah, 2002; Galon et al., 2008; Du et al., 2009 |
| er1 (Nt) | |||||
| cat2 | At4g35090 | Dismutation of H2O2 | Necrotic lesions in Long-Days conditions | Intracellular oxidative stress coupled with decreased of Myo-inositol content | Vandenabeele et al., 2004; Queval et al., 2007; Chaouch and Noctor, 2010 |
| cea62 (Os) | Os02g0110200 | hydroperoxyde lyase | Brown lesion spots over the entire leaf surface | Probably due to constitutive induction of JA signaling | Liu et al., 2012 |
| chs2 (gof) | At4g16860 | Encodes the TIR-NBS-LRR protein named RPP4 | Lesions and bleaching when transfer in low temperature conditions | Conditional activation of defense responses | Huang et al., 2010 |
| cpn1/bon1 | At5g61900 | Calcium-dependent phospholipid binding protein | Lesions appear on leaves in low humidity and/or low temperature conditions | Overexpression of SNC1 is necessary to induce lesions | Jambunathan et al., 2001; Jambunathan and McNellis, 2003; Yang and Hua, 2004 |
| cpr5 | At5g64930 | Transmembrane protein with unknown molecular function | Spontaneous lesions and precocious senescence | Unknown | Bowling et al., 1997; Kirik et al., 2001; Yoshida et al., 2002 |
| cpr22 (gof) | At2g46450 and At2g46440 | Cyclic-gated ion channel for Ca2+ | Chlorotic lesions, then plants die within 14 days after planting | The chimeric protein CNGC11/12 created by cpr22 deletion constitutively activate defense signaling | Yoshioka et al., 2001, 2006; Urquhart et al., 2007 |
| cslf6 (Os) | Os08g06380 | Cellulose synthase-like involved in mixed-linkage glucan biosynthesis | Spontaneous, discrete, necrotic lesions in flag and old leaves | Probably due to high decrease in mixed-linkage glucan content | Vega-Sanchez et al., 2012 |
| dnd1 | At5g15410 | Cyclic-gated ion channel for Ca2+ | Small necrotic lesions but also a decrease in HR induction | Unknown | Yu et al., 1998; Clough et al., 2000; Ali et al., 2007 |
| dnd2/hlm1 | At5g54250 | Cyclic-gated ion channel for K+, Na+ and/or Ca2+ | Spontaneus lesions but also a decrease in HR induction | Unknown | Balague et al., 2003; Jurkowski et al., 2004; Rostoks et al., 2006 |
| nec1 (Hv) | |||||
| edr1 | At1g08720 | Map Kinase Kinase Kinase | Necrotic lesions in the case of infection with the powdery mildew | Derepression of a MAP kinase cascade in edr1 seems to be responsible of the cell death phenotype | Frye and Innes, 1998; Frye et al., 2001; Zhao et al., 2014 |
| edr2 | At4g19040 | a PH and STAR domain containing protein | Necrotic lesions in the case of infection with the powdery mildew | Unknown | Tang et al., 2005 |
| edr3 | At3g60190 | Dynamin-related protein with a putative role in Golgi traffic | Necrotic lesions in the case of infection with the powdery mildew | Unknown | Hong et al., 2003; Tang et al., 2006 |
| erh1 | At2g37940 | Inositol phosphorylceramide synthase | Spontaneous lesions in a Col-0 genetic background expressing the resistance gene RPW8 | Ceramide accumulation | Wang et al., 2008 |
| erh1 (Os) | |||||
| exo70b1 | At5g58430 | Exocyst subunit EXO70 family protein B1 with a role in autophagy | HR-like lesions | Unknown | Kulich et al., 2013; Stegmann et al., 2013 |
| flu | At3g14110 | Protein with Coiled-coil TPR domain regulating chlorophyll biosynthesis | Lesions on mature leaves, complete bleaching of seedling after a dark to light transfer | Accumulation of Pchlide in the dark which then generate singlet oxygen in the light condition | Meskauskiene et al., 2001; Op Den Camp et al., 2003; Przybyla et al., 2008; Khandal et al., 2009 |
| tigrina-d.12 (Hv) | |||||
| fzl | At1g03160 | Membrane remodeling GTPase localized in chloroplast enveloppe | Chlorotic lesions on rosettes leaves in Ler ecotype but not in Col-0 | Probably caused by multiple chloroplasts anomalies observed in fzl | Gao et al., 2006; Landoni et al., 2013 |
| len1 | At1g55490 | Chaperonin 60β localized in chloroplast | Spontaneous lesions only in Short-days but not in Long-days condtions | Unknown | Ishikawa et al., 2003; Ishikawa, 2005 |
| les22 (Zm) | GRMZm2G044074 | Uroporphyrynogen decarboxylase (Chlorophyll biosynthesis) | HR-like necrotic lesions | Probably due to accumulation of uroporphyrine | Hu et al., 1998 |
| loh1 | At3g25540 | Ceramide synthase | lesions after a long growth in Short-days condition | Accumulation of free trihydroxy sphingoid bases or ceramide species with C16 fatty acids | Ternes et al., 2011 |
| lrgB | At1g32080 | Intermembrane protein localized in chloroplast envelop inner membrane | Chlorotic lesions on true leaves of juveniles plants | Unknown | Yamaguchi et al., 2012; Yang et al., 2012 |
| lin2 | At1g03475 | Coproporphyrinogen III oxidase involved in chlorophyll biosynthesis | Spontaneous lesions formation on young leaves | Probable accumulation of photosensitizing tetrapyrrole intermediates | Ishikawa et al., 2001; Sun et al., 2011; Guo et al., 2013 |
| Rlin1 (Os) | |||||
| lms (Os) | Os02g0639000 | Protein with a CTD phosphatase domain and two dsRBM motifs | Reddish-brown lesions on leaves and rapid senescence after flowering | Unknown | Undan et al., 2012 |
| lsd1 | At4g20380 | Protein containing three zinc finger domains | Spontaneous and runaway cell death on leaves when plants are transferred from Short-days to Long-days conditions or submitted to a highlight stress | Uncontrolled oxidative stress (anion superoxide and oxydized plastoquinone pool) that is the consequence of misregulation of numerous genes | Jabs et al., 1996; Dietrich et al., 1997; Kliebenstein et al., 1999; Muhlenbock et al., 2008 |
| lsd1 (Os) | |||||
| mips1 | At4g39800 | Myo-inositol-1-phosphate synthase which catalyze the first reaction of Myo-inositol biosynthesis | Necrotic lesions appear on rosette leaves after a transfer from Short-days to Long-days conditions | Not clearly determined but a ceramide accumulation can be observed | Meng et al., 2009; Donahue et al., 2010; Luo et al., 2011 |
| mkk1/mkk2 | At4g26070, At4g29810 | Two Map kinase kinase which act to negatively regulate plant defenses | Reduced growth, lesions that engulfed the whole seedling and lead to lethality | Constitutive activation of defense responses | Ichimura et al., 1998; Gao et al., 2008; Qiu et al., 2008 |
| mpk4 | At4g01370 | Map kinase which acts downstream to MKK1/MKK2 to negatively regulate plant defenses | Reduced growth, small spotted lesions | Constitutive activation of defense responses | Petersen et al., 2000; Su et al., 2007; Gao et al., 2008; Liu et al., 2011 |
| mpk4 (Gm) | |||||
| nsl1 | At1g28380 | Protein containing a MACPF domain | Spotted necrotic lesions | Unknown | Noutoshi et al., 2006 |
| pub13 | At3g46510, Os12g0570000 | Protein with E3 ubiquitin ligase activity | Chlorosis and spotted lesions on leaves in LD conditions | Constitutive activation of defense responses | Zeng et al., 2004; Li et al., 2012 |
| spl11 (Os) | |||||
| rug1 | At5g08280 | Porphobilinogen deaminase involved in chlorophyll biosynthesis | Spontaneous lesions on leaves | Probably due to accumulation of porphobilinogen | Quesada et al., 2013 |
| siz1 | At5g60410 | SUMO E3 ligase | Spontaneous lesions on leaves and dwarfism | Constitutive activation of defense responses | Lee et al., 2007a; Miura et al., 2010 |
| slh1 | At5g45260 | TIR-NBS-LRR with a WRKY domain | Necrotic lesions on rosette and cauline leaves in low humidity | Defect in DNA binding of the protein that conduces to Constitutive activation of defense responses | Noutoshi et al., 2005 |
| smg7 | At5g19400 | Involved in Nonsense-Mediated RNA decay process | Necrotic lesions on rosette leaves and lethality (depending on allele) | Constitutive activation of defense responses | Riehs-Kearnan et al., 2012 |
| spl7 (Os) | Os05g0530400 | Heat stress transcription factor | Small, reddish brown lesions over the whole surface of leaves | Unknown | Yamanouchi et al., 2002 |
| spl28 (Os) | Os01g0703600 | Clathrin adaptator subunit (AP1M1) involved in post-golgi traffic | Spotted lesions on leaves | Unknown | Qiao et al., 2010 |
| ssi2 | At2g43710 | Steroyl-ACP desaturase involved in fatty acid desaturation | HR-like lesions | Increase in 18:0 and decrease in 18:1 fatty acids, inducing, among other things, NO accumulation | Kachroo et al., 2001, 2008; Shah et al., 2001; Mandal et al., 2012 |
| ssi2 (Gm, Os) | |||||
| ssi4 (gof) | Putative TIR-NBS-LRR protein | Chlorotic lesions suppressed by high humidity | Probably caused by constitutive activation of defense signaling | Shirano et al., 2002; Zhou et al., 2004 | |
| syp121/syp122 | At3g11820, At3g52400 | Syntaxins involved in vesicular trafficking | Dwarf phenotype and necrotic lesions | Constitutive activation of defense responses | Zhang et al., 2008 |
| upf1 | At5g47010 | Involved in Nonsense-Mediated RNA decay process | Necrotic lesions on rosette leaves and lethality (depending on allele) | Constitutive activation of defense responses | Riehs-Kearnan et al., 2012 |
| vad1 | At1g02120 | Membrane-associated protein with a GRAM domain | Vascular necrotic lesions | Unknown | Lorrain et al., 2004 |
| xth27 | At2g01850 | Xyloglucan endotransglucosylase/hydrolase | Yellow necrotic spots associated with leaf maturation | Unknown | Matsui et al., 2005 |
Bo, Brassica oleracea; Gm, Glycine max; Hv, Hordeum vulgare (barley); Ta, Triticum aestivum; Zm, Zea mays; Nt, Nicotiana tabbacum; gof, the mutation in correspondings lmm is a Gain of function. In all other lmm, the mutation is recessive.