{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2025,10,12]],"date-time":"2025-10-12T03:26:42Z","timestamp":1760239602379,"version":"build-2065373602"},"reference-count":49,"publisher":"MDPI AG","issue":"24","license":[{"start":{"date-parts":[[2020,12,14]],"date-time":"2020-12-14T00:00:00Z","timestamp":1607904000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Molecules"],"abstract":"<jats:p>A stable intense resistance called \u201cnonhost resistance\u201d generates a complete multiple-gene resistance against plant pathogenic species that are not pathogens of pea such as the bean pathogen, Fusarium solani f. sp. phaseoli (Fsph). Chitosan is a natural nonhost resistance response gene activator of defense responses in peas. Chitosan may share with cancer-treatment compounds, netropsin and some anti-cancer drugs, a DNA minor groove target in plant host tissue. The chitosan heptamer and netropsin have the appropriate size and charge to reside in the DNA minor groove. The localization of a percentage of administered radio-labeled chitosan in the nucleus of plant tissue in vivo indicates its potential to transport to site(s) within the nuclear chromatin (1,2). Other minor groove-localizing compounds administered to pea tissue activate the same secondary plant pathway that terminates in the production of the anti-fungal isoflavonoid, pisatin an indicator of the generated resistance response. Some DNA minor groove compounds also induce defense genes designated as \u201cpathogenesis-related\u201d (PR) genes. Hypothetically, DNA targeting components alter host DNA in a manner enabling the transcription of defense genes previously silenced or minimally expressed. Defense-response-elicitors can directly (a) target host DNA at the site of transcription or (b) act by a series of cascading events beginning at the cell membrane and indirectly influence transcription. A single defense response, pisatin induction, induced by chitosan and compounds with known DNA minor groove attachment potential was followed herein. A hypothesis is formulated suggesting that this DNA target may be accountable for a portion of the defense response generated in nonhost resistance.<\/jats:p>","DOI":"10.3390\/molecules25245913","type":"journal-article","created":{"date-parts":[[2020,12,14]],"date-time":"2020-12-14T22:30:52Z","timestamp":1607985052000},"page":"5913","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Nonhost Disease Resistance in Pea: Chitosan\u2019s Suggested Role in DNA Minor Groove Actions Relative to Phytoalexin-Eliciting Anti-Cancer Compounds"],"prefix":"10.3390","volume":"25","author":[{"given":"Lee A.","family":"Hadwiger","sequence":"first","affiliation":[{"name":"Department of Plant Pathology, Washington State University, Pullman, WA 99164-6430, USA"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1968","published-online":{"date-parts":[[2020,12,14]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"170","DOI":"10.1104\/pp.67.1.170","article-title":"Localization of fungal components in the Pea-Fusarium interaction detected immunochemically with anti-chitosan and anti-fungal cell wall antisera","volume":"57","author":"Hadwiger","year":"1981","journal-title":"Plant Physiol."},{"key":"ref_2","unstructured":"Skjat-Braek, G., and Sandford, P.A. (1989). The molecular biology of chitosan in plant\/pathogen interaction and its application in agriculture. Chitin and Chitosan, Sources, Chemistry, Biochemistry, Physical Properties and Applications, Elsevier Applied Science."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"439","DOI":"10.1016\/j.plantsci.2009.07.007","article-title":"The non-host disease resistance in peas; alterations in phosphorylation and ubiquitination of HMGA and histone H2A\/H2B","volume":"177","author":"Isaac","year":"2009","journal-title":"Plant Sci."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"45","DOI":"10.1016\/j.plantsci.2006.07.014","article-title":"The use of chemical genomics to detect functional systems affecting the non-host disease resistance of pea to Fusarium solani f. sp.","volume":"172","author":"Hartney","year":"2007","journal-title":"phaseoli. Plant Sci."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"215","DOI":"10.1016\/0885-5765(89)90052-0","article-title":"Chitosan oligomers from Fusarium solani\/pea interactions, chitinase\/\u03b2-glucanase digestion of sporelings and from fungal wall chitin actively inhibit fungal growth and enhance disease resistance","volume":"35","author":"Kendra","year":"1989","journal-title":"Physiol. Mol. Plant Path."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"1056","DOI":"10.3389\/fpls.2018.01056","article-title":"DNA damage and chromatin conformation changes confer nonhost resistance: A hypothesis based on effects of anti-cancer agents on plant defense responses","volume":"9","author":"Hadwiger","year":"2018","journal-title":"Front. Plant Sci."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"373","DOI":"10.3389\/fpls.2015.00373","article-title":"Anatomy of a nonhost disease resistance response of pea to Fusarium solani: PR gene elicitation via DNase, chitosan and chromatin alterations","volume":"6","author":"Hadwiger","year":"2015","journal-title":"Front. Plant Sci."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"446","DOI":"10.3389\/fpls.2017.00446","article-title":"Nonhost resistance: DNA damage is associated with SA signaling for induction of PR genes and contributes to the growth suppression of a pea pathogen on pea endocarp tissue","volume":"8","author":"Hadwiger","year":"2017","journal-title":"Front. Plant Sci."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"81","DOI":"10.1094\/PHYTO-04-12-0085-R","article-title":"Fungal mitochondrial DNases. Effectors with the potential to activate plant defenses in nonhost resistance","volume":"1031","author":"Hadwiger","year":"2013","journal-title":"Phytopathology"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"18","DOI":"10.1016\/j.pmpp.2017.01.007","article-title":"Nonhost resistance: Reactive oxygen species (ROS) signal causes DNA damage and the induction of PR genes and disease resistance in pea tissue","volume":"98","author":"Tanaka","year":"2017","journal-title":"Physiol. Mol. Plant Pathol."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"278","DOI":"10.1006\/abbi.2000.1824","article-title":"Characterization of two novel defense peptides from pea (Pisum sativum) seeds","volume":"378","author":"Almeida","year":"2000","journal-title":"Arch. Biochem. Biophys."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"324","DOI":"10.1094\/MPMI-4-324","article-title":"The Fusarium solani-induced expression of a pea gene family encoding cysteine content proteins","volume":"4","author":"Chiang","year":"1991","journal-title":"Mol. Plant Microbe Interact."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"690","DOI":"10.1104\/pp.54.5.690","article-title":"Physiological and cytological similarities between disease resistance and cellular incompatibility responses","volume":"54","author":"Teasdale","year":"1974","journal-title":"Plant Physiol."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"1083","DOI":"10.1016\/j.ijbiomac.2018.01.113","article-title":"Identification of chitosan oligosaccharides binding proteins from the plasma membrane of wheat leaf cell","volume":"111","author":"Lui","year":"2018","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"607","DOI":"10.1104\/pp.76.3.607","article-title":"Ethylene: Symptom not signal for induction of chitinase and \u03b2-1, 3-glucanase in pea pods by pathogens and elicitors","volume":"76","author":"Mauch","year":"1984","journal-title":"Plant Physiol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"1376","DOI":"10.1073\/pnas.82.5.1376","article-title":"The molecular origin of DNA-drug specificity in netropsin and distamycin","volume":"82","author":"Kopka","year":"1985","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"4792","DOI":"10.1021\/bi9628228","article-title":"Crystal structure of the DNA decamer d(CGCAATTGCG) complexed with the minor groove binding drug netropsin","volume":"36","author":"Nunn","year":"1997","journal-title":"Biochemistry"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"1051","DOI":"10.3762\/bjoc.14.93","article-title":"An overview of recent advances in duplex DNA recognition by small molecules","volume":"14","author":"Bhaduri","year":"2018","journal-title":"Beilstein J. Org. Chem."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"9649","DOI":"10.1093\/nar\/gkr699","article-title":"Complexity in the binding of minor groove agents: Netropsin has two thermodynamically different DNA binding modes at a single site","volume":"39","author":"Lewis","year":"2011","journal-title":"Nucleic Acids Res."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"1248","DOI":"10.1038\/nature08473","article-title":"The Role of DNA shape in protein-DNA recognition","volume":"461","author":"Rohs","year":"2009","journal-title":"Nature"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"291","DOI":"10.1039\/a705982e","article-title":"DNA minor-groove recognition by small molecules","volume":"18","author":"Neidle","year":"2001","journal-title":"Nat. Prod. Rep."},{"key":"ref_22","first-page":"15","article-title":"Inverse PCR to identify DNA sequence upstream of the pea HMY I\/Y open reading frame","volume":"37","author":"Druffel","year":"2005","journal-title":"Pisum Genet."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"1312","DOI":"10.1094\/Phyto-74-1312","article-title":"Phytoalexin production in five cultivars of peas differentially resistance to three races of Pseudomonas syringe pv. pisi","volume":"74","author":"Hadwiger","year":"1984","journal-title":"Phytopathology"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"S1360","DOI":"10.1016\/S1360-1385(98)01199-6","article-title":"Pathogen defence in plants\u2014A paradigm of biological complexity","volume":"3","author":"Somssich","year":"1998","journal-title":"Trends Plant Sci."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"15","DOI":"10.1016\/0048-4059(85)90053-0","article-title":"Gene expression in susceptible and disease resistant interactions of Peas induced with Fusarium solani pathogens and chitosan","volume":"27","author":"Fristenski","year":"1985","journal-title":"Physiol. Plant Pathol."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"309","DOI":"10.1007\/BF00021310","article-title":"Pea genes associated with non-host disease to Fusarium are also active in race-specific disease resistance to Pseudomononas","volume":"8","author":"Daniels","year":"1987","journal-title":"Plant Mol. Biol."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"28","DOI":"10.1007\/s00122-002-0985-2","article-title":"Quantitive trait loci for partial resistance to Aphanomyces root rot in pea","volume":"106","author":"Muehlbauer","year":"2002","journal-title":"Theor. Appl. Genet."},{"key":"ref_28","first-page":"4","article-title":"How Plants Resist Disease-A focus on \u201cnonhost resistance\u201d of pea to a bean pathogen","volume":"2","author":"Hadwiger","year":"2018","journal-title":"Openaccess ebooks"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"79","DOI":"10.1016\/j.pmpp.2015.08.003","article-title":"Nonhost resistance: Self-inflicted DNA damage by fungal DNase accumulation is a major factor in terminating fungal growth in the pea-Fusarium solani f. sp. phaseoli interaction","volume":"92","author":"Hadwiger","year":"2015","journal-title":"Physiol. Mol. Plant Pathol."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"33","DOI":"10.1006\/pmpp.1993.1038","article-title":"Characterization of a plant stimulated nuclease from Fusarium solani","volume":"43","author":"Gerhold","year":"1993","journal-title":"Physiol. Mol. Plant Pathol."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"950","DOI":"10.3389\/fgene.2020.00950","article-title":"Identification of Fusarium solani f. sp. pisi (Fsp) responsive genes in Pisum sativum","volume":"11","author":"Sharpe","year":"2020","journal-title":"Front. Genet."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"85","DOI":"10.1006\/pmpp.1999.0213","article-title":"The families of pathogenesis-related proteins, their activities and comparative analysis of PR-1 type proteins","volume":"55","year":"1999","journal-title":"Physiol. Mol. Plant Pathol."},{"key":"ref_33","first-page":"336","article-title":"Studies on phytoalexins. V. The structure of pisatin from Pisum sativum L.","volume":"14","author":"Perrin","year":"1962","journal-title":"Aust. J. Biol. Sci."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"438","DOI":"10.1111\/j.1751-1097.1998.tb05224.x","article-title":"Photoactivated psoralens elicit defense genes and phytoalexins production in the pea plant","volume":"67","author":"Parsons","year":"1998","journal-title":"Photochem. Photobiol."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"653","DOI":"10.1016\/j.molcel.2008.02.014","article-title":"Histone ubiquitination triggering gene activity","volume":"29","author":"Weake","year":"2008","journal-title":"Mol. Cell"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"276","DOI":"10.1016\/0147-5975(84)90013-6","article-title":"Characterization of the smallest chitosan oligomer that is maximally antifungal to Fusarium solani and elicits pisatin formation in Pisum sativum","volume":"8","author":"Kendra","year":"1984","journal-title":"Exp. Mycol."},{"key":"ref_37","doi-asserted-by":"crossref","unstructured":"Paul, A., Guo, P., Boykin, D.W., and Wilson, W.D. (2019). A new generation of minor-groove-binding heterocyclic diamidines that recognize G-C base pairs in an AT sequence context. Molecules, 24.","DOI":"10.3390\/molecules24050946"},{"key":"ref_38","first-page":"1","article-title":"Binding to the DNA minor groove by heterocyclic dications: From AT-specific monomers to GC recognition with dimers","volume":"8","author":"Nanjunda","year":"2012","journal-title":"Corr. Protoc. Nucleic Acid Chem."},{"key":"ref_39","doi-asserted-by":"crossref","unstructured":"Yang, Y., Ronge, X., Lui, S., Qin, Y., Li, K., Yu, H., and Li, P. (2019). Immunostimulatory effects of chitooligosaccharides on RAW 264.7 mouse macrophages via regulation of the MAPK and PI3K\/Akt signaling pathways. Mar. Drugs, 17.","DOI":"10.3390\/md17010036"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"163","DOI":"10.1007\/BF01977526","article-title":"Changes in plant metabolism associated with phytoalexin production","volume":"74","author":"Hadwiger","year":"1968","journal-title":"Neth. J. Plant Path."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"285","DOI":"10.1016\/S0147-5975(79)80054-7","article-title":"The fungicidal effect of chitosan on fungi of varying cell wall composition","volume":"3","author":"Allan","year":"1979","journal-title":"Exp. Mycol."},{"key":"ref_42","doi-asserted-by":"crossref","unstructured":"Matica, A.M., Aachmann, F.L., Tondervik, A., Sletta, H., and Ostafe, V. (2019). Chitosan as a wound dressing starting material: Antimicrobial properties and mode of action. Int. J. Mol. Sci., 20.","DOI":"10.3390\/ijms20235889"},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"522","DOI":"10.3389\/fpls.2016.00522","article-title":"Chitin oligosaccharide and chitosan oligosaccharide: Two similar but different plant elicitors","volume":"7","author":"Yin","year":"2016","journal-title":"Front. Plant Sci."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"19613","DOI":"10.1073\/pnas.0705147104","article-title":"CERK1, a LysM receptor kinase, is essential for chitin elicitor signaling in Arabidopsis","volume":"104","author":"Miya","year":"2007","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_45","first-page":"13","article-title":"HMG A proteins: Flexible regulators of transcription and chromatin structure","volume":"1519","author":"Reeves","year":"2001","journal-title":"Biochem. Biophys. Acta"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"249","DOI":"10.1046\/j.1364-3703.2003.00171.x","article-title":"Analysis of pea HMG I\/Y suggests a role in defense gene regulation","volume":"4","author":"Klosterman","year":"2003","journal-title":"Mol. Plant Pathol."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"105","DOI":"10.1146\/annurev.biophys.27.1.105","article-title":"Minor groove-binding architectural proteins: Structure, function and RNA recognition","volume":"27","author":"Bewley","year":"1998","journal-title":"Ann. Rev. Biophys. Biomed. Struct."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"5550","DOI":"10.1093\/nar\/gkp558","article-title":"Large, sequence-dependent effects on DNA conformation by minor groove binding compounds","volume":"37","author":"Tavis","year":"2009","journal-title":"Nucleic Acids Res."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"1580","DOI":"10.1126\/science.1235441","article-title":"The supercoiling of DNA is a removable barrier to the RNA polymerase complex (RNAP) transcription of genes","volume":"340","author":"Ma","year":"2013","journal-title":"Science"}],"container-title":["Molecules"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1420-3049\/25\/24\/5913\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T10:44:40Z","timestamp":1760179480000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1420-3049\/25\/24\/5913"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2020,12,14]]},"references-count":49,"journal-issue":{"issue":"24","published-online":{"date-parts":[[2020,12]]}},"alternative-id":["molecules25245913"],"URL":"https:\/\/doi.org\/10.3390\/molecules25245913","relation":{},"ISSN":["1420-3049"],"issn-type":[{"type":"electronic","value":"1420-3049"}],"subject":[],"published":{"date-parts":[[2020,12,14]]}}}