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Penicillium verrucosum
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<h2 id="taxonomy">Taxonomy</h2> <p><i>Penicillium verrucosum</i> was initially incorrectly placed in synonymy with the species <i>Penicillium viridicatum</i> by Raper and Thom.<a class="footnote-ref" id="fnref:7" href="#fn:7"><sup>7</sup></a> Later, after disagreements arose upon the identification and naming of these fungi, their growth rates, mycotoxin productions and sources were observed.<a class="footnote-ref" id="fnref:8" href="#fn:8"><sup>8</sup></a> Careful observation concluded that <i>P. verrucosum</i> and <i>P. viridicatum</i> were indeed separate species. Experimental results showed that the mycotoxins <a href="/facts/Ochratoxin_A/yopAhdob">ochratoxin A</a> and <a href="/facts/Citrinin/Ve1gdu6b">citrinin</a> are produced by <i>P. verrucosum</i> but not by <i>P. viridicatum</i>.<a class="footnote-ref" id="fnref:9" href="#fn:9"><sup>9</sup></a> </p> <h2 id="habitat-and-ecology">Habitat and ecology</h2> <p><i>Penicillium verrucosum</i> is found in temperate and cooler climates.<a class="footnote-ref" id="fnref:10" href="#fn:10"><sup>10</sup></a> It is found predominantly in northern Europe, including countries such as Scandinavia, Ukraine, Denmark, Sweden, United Kingdom, Yugoslavia, Italy, and Turkey,<a class="footnote-ref" id="fnref:11" href="#fn:11"><sup>11</sup></a><a class="footnote-ref" id="fnref:12" href="#fn:12"><sup>12</sup></a> certain parts of North America, such as Canada,<a class="footnote-ref" id="fnref:13" href="#fn:13"><sup>13</sup></a> and parts of South America.<a class="footnote-ref" id="fnref:14" href="#fn:14"><sup>14</sup></a> <i>P. verrucosum</i> grows on grains, seeds and decaying vegetation.<a class="footnote-ref" id="fnref:15" href="#fn:15"><sup>15</sup></a> It is widely distributed in food (such as cereal<a class="footnote-ref" id="fnref:16" href="#fn:16"><sup>16</sup></a>) and animal feed where grains (usually barley, wheat and rye<a class="footnote-ref" id="fnref:17" href="#fn:17"><sup>17</sup></a>) are a key ingredient.<a class="footnote-ref" id="fnref:18" href="#fn:18"><sup>18</sup></a> </p> <h2 id="growth-and-morphology">Growth and morphology</h2> <p><i>Penicillium verrucosum</i> is found to be slow-growing: it achieves between 15 mm and 25 mm of growth in diameter on both <a href="/facts/Czapek_medium/3tqruVlT">Czapek Yeast Agar</a> (CYA) and Malt Extract Agar (MEA) after seven days.<a class="footnote-ref" id="fnref:19" href="#fn:19"><sup>19</sup></a> <i>P. verrucosum</i> has a white <a href="/facts/Mycelium/znB97XNn">mycelium</a> and greyish-green to dull green <a href="/facts/Conidium/klClhD20">conidia</a> on the aforementioned media.<a class="footnote-ref" id="fnref:20" href="#fn:20"><sup>20</sup></a> The reverse is coloured yellow brown to deep brown on CYA and dull brown to olive on MEA.<a class="footnote-ref" id="fnref:21" href="#fn:21"><sup>21</sup></a> Other varieties of <i>P. verrucosum</i> can have differently coloured conidia, including the colours dark green and blue-green.<a class="footnote-ref" id="fnref:22" href="#fn:22"><sup>22</sup></a> This fungus has a greater number of conidia growth on CYA than MEA.<a class="footnote-ref" id="fnref:23" href="#fn:23"><sup>23</sup></a> The conidia are smooth-walled and approximately 2.5 μm to 3.0 μm in diameter.<a class="footnote-ref" id="fnref:24" href="#fn:24"><sup>24</sup></a> These conidia begin in an ellipsoidal shape when young, and later change to a globose or subglobose shape.<a class="footnote-ref" id="fnref:25" href="#fn:25"><sup>25</sup></a> <i>P. verrucosum</i> possesses conidiophores which are usually two-stage branched (sometimes three-stage branched), giving it a brush-like appearance.<a class="footnote-ref" id="fnref:26" href="#fn:26"><sup>26</sup></a> The conidiophores of <i>P. verrucosum</i> are rough-walled with branches and metulae that are pressed closely together.<a class="footnote-ref" id="fnref:27" href="#fn:27"><sup>27</sup></a> The <a href="/facts/Phialide/TIvs3RwC">phialides</a> of the conidiophore are short and flask-shaped with distinct necks.<a class="footnote-ref" id="fnref:28" href="#fn:28"><sup>28</sup></a> </p><p><i>Penicillium verrucosum</i> has a distinctive odour which is described as earthy and pungent.<a class="footnote-ref" id="fnref:29" href="#fn:29"><sup>29</sup></a> </p> <h2 id="physiology">Physiology</h2> <p>Of the genus <i>Penicillium</i>, only about half of the identified species are able to grow at the body temperature of mammals.<a class="footnote-ref" id="fnref:30" href="#fn:30"><sup>30</sup></a> <i>P. verrucosum</i> is not one of these species as there is usually no growth of this fungus at 37 °C.<a class="footnote-ref" id="fnref:31" href="#fn:31"><sup>31</sup></a> The conidia of <i>P. verrucosum</i> have the ability to germinate at temperatures between 0 °C and 31 °C,<a class="footnote-ref" id="fnref:32" href="#fn:32"><sup>32</sup></a> but optimal temperatures for germination are between 21 °C and 23 °C.<a class="footnote-ref" id="fnref:33" href="#fn:33"><sup>33</sup></a> Metabolic products of this fungi include 2-octen-1-ol and <a href="/facts/1-Octanol/pWEETXd5">1-octanol</a> and ochratoxin A, <a href="/facts/Brevianamide_a_and_b/tjufPPgK">brevianamide A</a>, citrinin, <a href="/facts/Penicillic_acid/sm7l7lXI">penicillic acid</a>, <a href="/facts/Ergosterol/Xt5FV1ut">ergosterol</a>, ergosteryl palmitate, meso-erythritol, <a href="/facts/Mannitol/CeHJsKMq">mannitol</a>, viridicatic acid, viridicatol, viridicatin, xanthomegnin, viomellein, rubrosulphin, viopurpurin, 3-O-methylviridicatin, cyclopenin, cyclopenol.<a class="footnote-ref" id="fnref:34" href="#fn:34"><sup>34</sup></a> </p> <h3>Ochratoxin A</h3> <p><i>Penicillium verrucosum</i> produces a very potent mycotoxin called ochratoxin A (OTA).<a class="footnote-ref" id="fnref:35" href="#fn:35"><sup>35</sup></a> This mycotoxin is immunosuppressive and <a href="/facts/Teratogenic/j4BN6lsB">teratogenic</a>.<a class="footnote-ref" id="fnref:36" href="#fn:36"><sup>36</sup></a> It has also been classified as <a href="/facts/Genotoxicity/fRUJaqxo">genotoxic</a><a class="footnote-ref" id="fnref:37" href="#fn:37"><sup>37</sup></a> and a possible human <a href="/facts/Carcinogen/soH8Y9CE">carcinogen</a>.<a class="footnote-ref" id="fnref:38" href="#fn:38"><sup>38</sup></a> Pigs raised in northern and central Europe develop nephritis after consumption of contaminated feed.<a class="footnote-ref" id="fnref:39" href="#fn:39"><sup>39</sup></a> The consumption of contaminated barley has been found to be toxic to rats and of contaminated rice has proven to be toxic to mice.<a class="footnote-ref" id="fnref:40" href="#fn:40"><sup>40</sup></a> </p> <h4>Storage conditions</h4> <p><i>Penicillium verrucosum</i> is a contaminant of cereal crops (such as barley, maize, oats and wheat) which are used in animal feed.<a class="footnote-ref" id="fnref:41" href="#fn:41"><sup>41</sup></a> These grains become contaminated with this mycotoxin when they are not carefully prepared after harvest and when storage conditions are unsuitable.<a class="footnote-ref" id="fnref:42" href="#fn:42"><sup>42</sup></a> When grains are properly stored, OTA levels tend to average around 1 μg/kg in temperate areas.<a class="footnote-ref" id="fnref:43" href="#fn:43"><sup>43</sup></a> </p><p>Spoilage of the grain by OTA occurs as a cause of inappropriate storage temperatures and moisture content.<a class="footnote-ref" id="fnref:44" href="#fn:44"><sup>44</sup></a> OTA synthesis occurs at moisture content levels between 18% and 22%, and OTA production increases when temperatures are between 10 °C and 21 °C.<a class="footnote-ref" id="fnref:45" href="#fn:45"><sup>45</sup></a> OTA formation does not occur at moisture content levels below 18% and at temperatures above 28 °C.<a class="footnote-ref" id="fnref:46" href="#fn:46"><sup>46</sup></a> </p> <h4>Prevention and reduction of contamination</h4> <p>To prevent OTA formation, grains must be dried to moisture levels lower than the 18% limit shortly after harvest.<a class="footnote-ref" id="fnref:47" href="#fn:47"><sup>47</sup></a> Any spoiled commodities should be kept apart from the uncontaminated harvest, and should not be used in food or feed production.<a class="footnote-ref" id="fnref:48" href="#fn:48"><sup>48</sup></a> Many countries have regulations regarding recommended and permitted levels of OTA in grains which should be followed.<a class="footnote-ref" id="fnref:49" href="#fn:49"><sup>49</sup></a> </p><p>Complete prevention of OTA contamination is ideal, but many methods exist to reduce existing OTA levels, usually classified into physical, chemical and biological procedures.<a class="footnote-ref" id="fnref:50" href="#fn:50"><sup>50</sup></a> Physical methods are used remove the contaminated grains through sorting and separation.<a class="footnote-ref" id="fnref:51" href="#fn:51"><sup>51</sup></a> Chemical procedures aim to eliminate this mycotoxin through processes such as <a href="/facts/Ammonium/jYm8wc1n">ammoniation</a>, <a href="/facts/Ozonation/o0E5gn2p">ozonation</a> and <a href="/facts/Nixtamalization/0LSETjgR">nixtamalization</a>.<a class="footnote-ref" id="fnref:52" href="#fn:52"><sup>52</sup></a> Biological processes use microorganisms to decompose or adsorb OTA in contaminated commodities.<a class="footnote-ref" id="fnref:53" href="#fn:53"><sup>53</sup></a> <a href="/facts/Protozoa/JczTf25H">Protozoa</a>, <a href="/facts/Bacteria/wC8xSCsU">bacteria</a>, <a href="/facts/Yeast/eGD24nfI">yeast</a>, <a href="/facts/Filamentous_fungus/7nHqptM7">filamentous fungi</a> and plant cell cultures are all used in these biological procedures.<a class="footnote-ref" id="fnref:54" href="#fn:54"><sup>54</sup></a> Microorganisms are beneficial for this purpose as they are environmentally-friendly and do not affect the grain quality.<a class="footnote-ref" id="fnref:55" href="#fn:55"><sup>55</sup></a> </p> <h2 id="disease-in-humans">Disease in humans</h2> <p>During the 1950s, reports of kidney disease with high rates of mortality were occurring in geographically close areas, such as Bulgaria, Yugoslavia and Romania.<a class="footnote-ref" id="fnref:56" href="#fn:56"><sup>56</sup></a> This occurrence was called the Balkan Endemic Nephropathy<a class="footnote-ref" id="fnref:57" href="#fn:57"><sup>57</sup></a> that was being caused by the consumption of contaminated pig meat in those areas.<a class="footnote-ref" id="fnref:58" href="#fn:58"><sup>58</sup></a> When pigs consumed feed contaminated by OTA, it collected in their fatty tissue rather than being excreted due to its solubility in fat.<a class="footnote-ref" id="fnref:59" href="#fn:59"><sup>59</sup></a> Humans then consumed contaminated pig meat, allowing this mycotoxin to enter the human system.<a class="footnote-ref" id="fnref:60" href="#fn:60"><sup>60</sup></a> </p><p>OTA is mainly found in blood samples in Europe,<a class="footnote-ref" id="fnref:61" href="#fn:61"><sup>61</sup></a> but its presence in healthy human blood shows that there is still worldwide exposure.<a class="footnote-ref" id="fnref:62" href="#fn:62"><sup>62</sup></a> An effort has been made in Europe to monitor the OTA levels in foods by creating regulations regarding maximum acceptable levels.<a class="footnote-ref" id="fnref:63" href="#fn:63"><sup>63</sup></a> Creating guidelines allows for special attention to be paid to local specialties, such as blood puddings and sausages, which are made using pig blood.<a class="footnote-ref" id="fnref:64" href="#fn:64"><sup>64</sup></a> </p> <h2 id="references">References</h2> <ol> <li id="fn:1"><p>Stoll, D.A.; Geisen, R.; Schmidt-Heydt, M. "Whole Genome Shotgun Sequencing of Penicillium verrucosum reveals about 42 secondary metabolite gene clusters". {{cite web}}: Missing or empty |url= (help) <a href="/wiki/Template:Cite_web" target="_blank">/wiki/Template:Cite_web</a> <a href="#fnref:1" class="footnote-back-ref">↩</a></p></li> <li id="fn:2"><p>"Penicillium verrucosum". MycoBank Database. Retrieved 7 October 2016. <a href="http://www.mycobank.org/BioloMICS.aspx?Table=Mycobank&Rec=37124&Fields=All" target="_blank">http://www.mycobank.org/BioloMICS.aspx?Table=Mycobank&Rec=37124&Fields=All</a> <a href="#fnref:2" class="footnote-back-ref">↩</a></p></li> <li id="fn:3"><p>"Penicillium verrucosum Dierckx". MycoBank Database. Retrieved 13 October 2016. <a href="http://www.mycobank.org/BioloMICS.aspx?TableKey=14682616000000063&Rec=14971&Fields=All" target="_blank">http://www.mycobank.org/BioloMICS.aspx?TableKey=14682616000000063&Rec=14971&Fields=All</a> <a href="#fnref:3" class="footnote-back-ref">↩</a></p></li> <li id="fn:4"><p>"Penicillium verrucosum Dierckx". MycoBank Database. Retrieved 13 October 2016. <a href="http://www.mycobank.org/BioloMICS.aspx?TableKey=14682616000000063&Rec=14971&Fields=All" target="_blank">http://www.mycobank.org/BioloMICS.aspx?TableKey=14682616000000063&Rec=14971&Fields=All</a> <a href="#fnref:4" class="footnote-back-ref">↩</a></p></li> <li id="fn:5"><p>Cabanes, Francisco Javier; Bragulat, Maria Rosa; Castella, Gemma (2010). "Ochratoxin A producing species in the Genus Penicillium". Toxins. 2 (5): 1111–1120. doi:10.3390/toxins2051111. PMC 3153233. PMID 22069629. <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3153233" target="_blank">https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3153233</a> <a href="#fnref:5" class="footnote-back-ref">↩</a></p></li> <li id="fn:6"><p>Stoll, D.A.; Geisen, R.; Schmidt-Heydt, M. "Whole Genome Shotgun Sequencing of Penicillium verrucosum reveals about 42 secondary metabolite gene clusters". {{cite web}}: Missing or empty |url= (help) <a href="/wiki/Template:Cite_web" target="_blank">/wiki/Template:Cite_web</a> <a href="#fnref:6" class="footnote-back-ref">↩</a></p></li> <li id="fn:7"><p>Raper, K.B.; Thom, C. (1949). A manual of the penicillia. Baltimore: The Williams & Wilkins Co. <a href="https://archive.org/details/in.ernet.dli.2015.549218" target="_blank">https://archive.org/details/in.ernet.dli.2015.549218</a> <a href="#fnref:7" class="footnote-back-ref">↩</a></p></li> <li id="fn:8"><p>Pitt, John (1987). "Penicillium viridicatum, Penicillium verrucosum, and production of ochratoxin A". Applied and Environmental Microbiology. 53 (2): 266–269. Bibcode:1987ApEnM..53..266P. doi:10.1128/aem.53.2.266-269.1987. PMC 203649. PMID 3566267. <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC203649" target="_blank">https://www.ncbi.nlm.nih.gov/pmc/articles/PMC203649</a> <a href="#fnref:8" class="footnote-back-ref">↩</a></p></li> <li id="fn:9"><p>Pitt, John (1987). "Penicillium viridicatum, Penicillium verrucosum, and production of ochratoxin A". Applied and Environmental Microbiology. 53 (2): 266–269. Bibcode:1987ApEnM..53..266P. doi:10.1128/aem.53.2.266-269.1987. PMC 203649. PMID 3566267. <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC203649" target="_blank">https://www.ncbi.nlm.nih.gov/pmc/articles/PMC203649</a> <a href="#fnref:9" class="footnote-back-ref">↩</a></p></li> <li id="fn:10"><p>International Society for Human and Animal Mycology (1994). Journal of Medical and Veterinary Mycology, Volume 32, Supplement 1. Oxford, UK: Blackwell Scientific Publications. <a href="#fnref:10" class="footnote-back-ref">↩</a></p></li> <li id="fn:11"><p>International Society for Human and Animal Mycology (1994). Journal of Medical and Veterinary Mycology, Volume 32, Supplement 1. Oxford, UK: Blackwell Scientific Publications. <a href="#fnref:11" class="footnote-back-ref">↩</a></p></li> <li id="fn:12"><p>Domsch, K.H.; Gams, Walter; Andersen, Traute-Heidi (1980). Compendium of soil fungi (2nd ed.). London, UK: Academic Press. ISBN 9780122204029. <a href="9780122204029" target="_blank">9780122204029</a> <a href="#fnref:12" class="footnote-back-ref">↩</a></p></li> <li id="fn:13"><p>Kozakiewicz, Z. "Penicillium verrucosum". CABI Bioscience. Retrieved 13 October 2016. <a href="http://www.cabi.org/dfb/abstract/20056401550" target="_blank">http://www.cabi.org/dfb/abstract/20056401550</a> <a href="#fnref:13" class="footnote-back-ref">↩</a></p></li> <li id="fn:14"><p>Lawley, Richard. "Ochratoxins". Food Safety Watch. Retrieved 7 November 2016. <a href="http://www.foodsafetywatch.org/factsheets/ochratoxins/" target="_blank">http://www.foodsafetywatch.org/factsheets/ochratoxins/</a> <a href="#fnref:14" class="footnote-back-ref">↩</a></p></li> <li id="fn:15"><p>International Society for Human and Animal Mycology (1994). Journal of Medical and Veterinary Mycology, Volume 32, Supplement 1. Oxford, UK: Blackwell Scientific Publications. <a href="#fnref:15" class="footnote-back-ref">↩</a></p></li> <li id="fn:16"><p>Cabanes, Francisco Javier; Bragulat, Maria Rosa; Castella, Gemma (2010). "Ochratoxin A producing species in the Genus Penicillium". Toxins. 2 (5): 1111–1120. doi:10.3390/toxins2051111. PMC 3153233. PMID 22069629. <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3153233" target="_blank">https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3153233</a> <a href="#fnref:16" class="footnote-back-ref">↩</a></p></li> <li id="fn:17"><p>Cabanes, Francisco Javier; Bragulat, Maria Rosa; Castella, Gemma (2010). "Ochratoxin A producing species in the Genus Penicillium". Toxins. 2 (5): 1111–1120. doi:10.3390/toxins2051111. PMC 3153233. PMID 22069629. <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3153233" target="_blank">https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3153233</a> <a href="#fnref:17" class="footnote-back-ref">↩</a></p></li> <li id="fn:18"><p>Kozakiewicz, Z. "Penicillium verrucosum". CABI Bioscience. Retrieved 13 October 2016. <a href="http://www.cabi.org/dfb/abstract/20056401550" target="_blank">http://www.cabi.org/dfb/abstract/20056401550</a> <a href="#fnref:18" class="footnote-back-ref">↩</a></p></li> <li id="fn:19"><p>Pitt, J.I.; Hocking, A.D. (1999). Fungi and food spoilage (2nd ed.). Gaithersburg, Md.: Aspen Publications. ISBN 978-0834213067. <a href="978-0834213067" target="_blank">978-0834213067</a> <a href="#fnref:19" class="footnote-back-ref">↩</a></p></li> <li id="fn:20"><p>Pitt, J.I.; Hocking, A.D. (1999). Fungi and food spoilage (2nd ed.). Gaithersburg, Md.: Aspen Publications. ISBN 978-0834213067. <a href="978-0834213067" target="_blank">978-0834213067</a> <a href="#fnref:20" class="footnote-back-ref">↩</a></p></li> <li id="fn:21"><p>Pitt, J.I.; Hocking, A.D. (1999). Fungi and food spoilage (2nd ed.). Gaithersburg, Md.: Aspen Publications. ISBN 978-0834213067. <a href="978-0834213067" target="_blank">978-0834213067</a> <a href="#fnref:21" class="footnote-back-ref">↩</a></p></li> <li id="fn:22"><p>Samson, Robert A.; Hoekstra, Ellen S.; Frisvad, Jens C. (2004). Introduction to food- and airborne fungi (7th ed.). Washington, DC: ASM Press. ISBN 978-9070351526. <a href="978-9070351526" target="_blank">978-9070351526</a> <a href="#fnref:22" class="footnote-back-ref">↩</a></p></li> <li id="fn:23"><p>Pitt, J.I.; Hocking, A.D. (1999). Fungi and food spoilage (2nd ed.). Gaithersburg, Md.: Aspen Publications. ISBN 978-0834213067. <a href="978-0834213067" target="_blank">978-0834213067</a> <a href="#fnref:23" class="footnote-back-ref">↩</a></p></li> <li id="fn:24"><p>Pitt, J.I.; Hocking, A.D. (1999). Fungi and food spoilage (2nd ed.). Gaithersburg, Md.: Aspen Publications. ISBN 978-0834213067. <a href="978-0834213067" target="_blank">978-0834213067</a> <a href="#fnref:24" class="footnote-back-ref">↩</a></p></li> <li id="fn:25"><p>Domsch, K.H.; Gams, Walter; Andersen, Traute-Heidi (1980). Compendium of soil fungi (2nd ed.). London, UK: Academic Press. ISBN 9780122204029. <a href="9780122204029" target="_blank">9780122204029</a> <a href="#fnref:25" class="footnote-back-ref">↩</a></p></li> <li id="fn:26"><p>Rippon, John Willard (1988). Medical mycology: the pathogenic fungi and the pathogenic actinomycetes (3rd ed.). Philadelphia, PA: Saunders. ISBN 978-0721624440. <a href="978-0721624440" target="_blank">978-0721624440</a> <a href="#fnref:26" class="footnote-back-ref">↩</a></p></li> <li id="fn:27"><p>Samson, Robert A.; Hoekstra, Ellen S.; Frisvad, Jens C. (2004). Introduction to food- and airborne fungi (7th ed.). Washington, DC: ASM Press. ISBN 978-9070351526. <a href="978-9070351526" target="_blank">978-9070351526</a> <a href="#fnref:27" class="footnote-back-ref">↩</a></p></li> <li id="fn:28"><p>Samson, Robert A.; Hoekstra, Ellen S.; Frisvad, Jens C. (2004). Introduction to food- and airborne fungi (7th ed.). Washington, DC: ASM Press. 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