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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Nutr.</journal-id>
<journal-title>Frontiers in Nutrition</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Nutr.</abbrev-journal-title>
<issn pub-type="epub">2296-861X</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
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<article-meta>
<article-id pub-id-type="doi">10.3389/fnut.2024.1384273</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Nutrition</subject>
<subj-group>
<subject>Perspective</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>Mushrooms: a food-based solution to vitamin D deficiency to include in dietary guidelines</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes">
<name><surname>Starck</surname> <given-names>Carlene</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="corresp" rid="c001"><sup>&#x002A;</sup></xref>
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<contrib contrib-type="author">
<name><surname>Cassettari</surname> <given-names>Tim</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
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<contrib contrib-type="author">
<name><surname>Wright</surname> <given-names>Jutta</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
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<contrib contrib-type="author">
<name><surname>Petocz</surname> <given-names>Peter</given-names></name>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
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<contrib contrib-type="author">
<name><surname>Beckett</surname> <given-names>Emma</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="aff" rid="aff3"><sup>3</sup></xref>
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<contrib contrib-type="author" corresp="yes">
<name><surname>Fayet-Moore</surname> <given-names>Flavia</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="aff" rid="aff4"><sup>4</sup></xref>
<xref ref-type="corresp" rid="c002"><sup>&#x002A;</sup></xref>
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<aff id="aff1"><sup>1</sup><institution>FOODiQ Global</institution>, <addr-line>Sydney, NSW</addr-line>, <country>Australia</country></aff>
<aff id="aff2"><sup>2</sup><institution>Macquarie University</institution>, <addr-line>Sydney, NSW</addr-line>, <country>Australia</country></aff>
<aff id="aff3"><sup>3</sup><institution>School of Health Sciences, The University of New South Wales</institution>, <addr-line>Kensington, NSW</addr-line>, <country>Australia</country></aff>
<aff id="aff4"><sup>4</sup><institution>School of Environmental and Life Sciences, The University of Newcastle</institution>, <addr-line>Callaghan, NSW</addr-line>, <country>Australia</country></aff>
<author-notes>
<fn fn-type="edited-by" id="fn0001"><p>Edited by: John Ayobami Amao, University of Ilorin, Nigeria</p></fn>
<fn fn-type="edited-by" id="fn0002"><p>Reviewed by: Gianluca Rizzo, Independent researcher, Messina, Italy</p></fn>
<corresp id="c001">&#x002A;Correspondence: Carlene Starck, <email>carlene@foodiq.global</email></corresp>
<corresp id="c002">Flavia Fayet-Moore, <email>flavia@foodiq.global</email></corresp>
</author-notes>
<pub-date pub-type="epub">
<day>10</day>
<month>04</month>
<year>2024</year>
</pub-date>
<pub-date pub-type="collection">
<year>2024</year>
</pub-date>
<volume>11</volume>
<elocation-id>1384273</elocation-id>
<history>
<date date-type="received">
<day>19</day>
<month>02</month>
<year>2024</year>
</date>
<date date-type="accepted">
<day>29</day>
<month>03</month>
<year>2024</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#x00A9; 2024 Starck, Cassettari, Wright, Petocz, Beckett and Fayet-Moore.</copyright-statement>
<copyright-year>2024</copyright-year>
<copyright-holder>Starck, Cassettari, Wright, Petocz, Beckett and Fayet-Moore</copyright-holder>
<license xlink:href="http://creativecommons.org/licenses/by/4.0/">
<p>This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.</p>
</license>
</permissions>
<abstract>
<p>Vitamin D deficiency and insufficiency is a public health issue, with low dietary vitamin D intakes a contributing factor. Rates of vitamin D deficiency are 31% in Australia, and up to 72% in some regions globally. While supplementation is often prescribed as an alternative to additional sun exposure, complementary approaches including food-based solutions are needed. Yet, food-centric dietary guidelines are not always adequate for meeting vitamin D needs. Edible mushrooms such as <italic>Agaricus bisporus</italic> can produce over 100% of vitamin D recommendations (10&#x2009;&#x03BC;g/day, Institute of Medicine) per 75&#x2009;g serve (18&#x2009;&#x03BC;g) on exposure to UV-light, with the vitamin D<sub>2</sub> produced showing good stability during cooking and processing. However, mushrooms are overlooked as a vitamin D source in dietary guidelines. Our dietary modelling shows that four serves/week of UV-exposed button mushrooms can support most Australian adults in meeting vitamin D recommendations, and UV-exposed mushrooms have been found to increase vitamin D status in deficient individuals. While recent evidence suggests some differences between vitamin D<sub>2</sub> and vitamin D<sub>3</sub> in physiological activities, vitamin D<sub>2</sub> from mushrooms can be part of a larger solution to increasing dietary vitamin D intakes, as well as an important focus for public health policy. Mushrooms exposed to UV represent an important tool in the strategic toolkit for addressing vitamin D deficiency in Australia and globally. Health authorities lead the recognition and promotion of mushrooms as a natural, vegan, safe, and sustainable vitamin D food source.</p>
</abstract>
<kwd-group>
<kwd>vitamin D deficiency</kwd>
<kwd>vitamin D intakes</kwd>
<kwd>mushrooms</kwd>
<kwd>dietary guidelines</kwd>
<kwd>recommended intake</kwd>
</kwd-group>
<counts>
<fig-count count="1"/>
<table-count count="1"/>
<equation-count count="0"/>
<ref-count count="71"/>
<page-count count="7"/>
<word-count count="6211"/>
</counts>
<custom-meta-wrap>
<custom-meta>
<meta-name>section-at-acceptance</meta-name>
<meta-value>Nutritional Epidemiology</meta-value>
</custom-meta>
</custom-meta-wrap>
</article-meta>
</front>
<body>
<sec id="sec1">
<title>Introduction: vitamin D deficiency is a global problem</title>
<p>Vitamin D deficiency is a global health concern with significant implications for population health. Systematic reviews (<xref ref-type="bibr" rid="ref1 ref2 ref3 ref4 ref5 ref6">1&#x2013;6</xref>) indicate that rates of vitamin D deficiency, when defined as blood level &#x003C;50&#x2009;nmol/L (<xref ref-type="bibr" rid="ref7">7</xref>, <xref ref-type="bibr" rid="ref8">8</xref>), are as high as 47.9% globally (range 19 to 72%) (<xref ref-type="table" rid="tab1">Table 1</xref>) (<xref ref-type="bibr" rid="ref1">1</xref>). In Australia, approximately three-quarters of the adult population have suboptimal vitamin D status; the rate of vitamin D deficiency is 31% (<xref ref-type="bibr" rid="ref1">1</xref>) and insufficiency (50 to 75&#x2009;nmol/L) a further 43% (<xref ref-type="bibr" rid="ref16">16</xref>). Some population groups are more vulnerable, with deficiency rates up to 94% in residential care-elderly (<xref ref-type="bibr" rid="ref17">17</xref>, <xref ref-type="bibr" rid="ref18">18</xref>). Along with a well-established role in bone health (<xref ref-type="bibr" rid="ref19">19</xref>), associations have been made between vitamin D inadequacy and increased susceptibility to infectious diseases (including COVID-19), muscle weakness, multiple sclerosis, diabetes, hypertension, metabolic syndrome, cancers, autoimmune diseases, cardiovascular disease (<xref ref-type="bibr" rid="ref1">1</xref>), and gestational diabetes (<xref ref-type="bibr" rid="ref20">20</xref>). While these associations are largely observational, the evidence is clear: the &#x201C;sunshine&#x201D; vitamin requires a metaphorical light to be shone upon it, and on a global scale.</p>
<table-wrap position="float" id="tab1">
<label>Table 1</label>
<caption>
<p>Rates of vitamin D deficiency and mean vitamin D intakes in Australia and World Health Organization global regions.</p>
</caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th align="left" valign="top">Country/region<xref ref-type="table-fn" rid="tfn3"><sup>c</sup></xref></th>
<th align="center" valign="top">Rate of vitamin D deficiency<xref ref-type="table-fn" rid="tfn1"><sup>a</sup></xref></th>
<th align="center" valign="top">Mean vitamin D intake (&#x03BC;g/day)<xref ref-type="table-fn" rid="tfn2"><sup>b</sup></xref></th>
<th align="center" valign="top">Additional references</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left" valign="top">Australia</td>
<td align="center" valign="top">31%</td>
<td align="center" valign="top">1.84 to 3.25</td>
<td align="center" valign="top">(<xref ref-type="bibr" rid="ref8">8</xref>)</td>
</tr>
<tr>
<td align="left" valign="top">Africas</td>
<td align="center" valign="top">19%</td>
<td align="center" valign="top">1 to 9.6</td>
<td align="center" valign="top">(<xref ref-type="bibr" rid="ref9">9</xref>)</td>
</tr>
<tr>
<td align="left" valign="top">Eastern Mediterranean</td>
<td align="center" valign="top">72%</td>
<td align="center" valign="top">1 to 4</td>
<td align="center" valign="top">(<xref ref-type="bibr" rid="ref10">10</xref>)</td>
</tr>
<tr>
<td align="left" valign="top">Europe</td>
<td align="center" valign="top">53%</td>
<td align="center" valign="top">2 to 4</td>
<td align="center" valign="top">(<xref ref-type="bibr" rid="ref10 ref11 ref12 ref13">10&#x2013;13</xref>)</td>
</tr>
<tr>
<td align="left" valign="top">Americas</td>
<td align="center" valign="top">30%</td>
<td align="center" valign="top">3.5 to 6</td>
<td align="center" valign="top">(<xref ref-type="bibr" rid="ref14">14</xref>, <xref ref-type="bibr" rid="ref15">15</xref>)</td>
</tr>
<tr>
<td align="left" valign="top">South-East Asia</td>
<td align="center" valign="top">57%</td>
<td align="center" valign="top">1.5 to 5.5</td>
<td align="center" valign="top">(<xref ref-type="bibr" rid="ref11">11</xref>)</td>
</tr>
<tr>
<td align="left" valign="top">Western Pacific</td>
<td align="center" valign="top">44%</td>
<td align="center" valign="top">1.84 to 7.6</td>
<td align="center" valign="top">(<xref ref-type="bibr" rid="ref8">8</xref>, <xref ref-type="bibr" rid="ref11">11</xref>)</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn id="tfn3">
<label>a</label>
<p>World Health Organisation (WHO) regions (<xref ref-type="bibr" rid="ref1">1</xref>), with Australia considered separately.</p>
</fn>
<fn id="tfn1">
<label>b</label>
<p>Defined as &#x003C;50&#x2009;nmol/L (<xref ref-type="bibr" rid="ref7">7</xref>, <xref ref-type="bibr" rid="ref8">8</xref>). Rates of deficiency sourced from Cui 2023 meta-analysis (<xref ref-type="bibr" rid="ref1">1</xref>).</p>
</fn>
<fn id="tfn2">
<label>c</label>
<p>Intake data sourced from the references listed in the final column of this table.</p>
</fn>
</table-wrap-foot>
</table-wrap>
<sec id="sec2">
<title>Food-based solutions can support vitamin D intake and status</title>
<p>Current vitamin D guidelines in Australia suggest that sunlight is a key source of vitamin D; where sunlight exposure is limited, supplements are the recommended alternative, with diet considered a poor source (<xref ref-type="bibr" rid="ref8">8</xref>, <xref ref-type="bibr" rid="ref18">18</xref>, <xref ref-type="bibr" rid="ref21 ref22 ref23">21&#x2013;23</xref>). Conversely, increased vitamin D intake from foods has consistently shown the ability to improve vitamin D status in those who have sub-optimal status (<xref ref-type="bibr" rid="ref24">24</xref>). Global vitamin D intakes are lower than the estimated average requirement (EAR) for vitamin D of 10&#x2009;&#x03BC;g/day set by the Institute of Medicine (IOM) (<xref ref-type="bibr" rid="ref25">25</xref>) (<xref ref-type="table" rid="tab1">Table 1</xref>) and recommended dietary patterns provided by dietary guidelines are insufficient for vitamin D (<xref ref-type="bibr" rid="ref21">21</xref>, <xref ref-type="bibr" rid="ref26">26</xref>).</p>
<p>Australian intakes of vitamin D are among the lowest in the world at 1.84 to 3.25&#x2009;&#x03BC;g/day (<xref ref-type="bibr" rid="ref8">8</xref>). While the potential for daily sunlight exposure is high in countries such as Australia, levels of exposure are insufficient to maintain vitamin D adequacy, with many factors suggested to play a role, including indoor lifestyles, skin color, and skin cancer risk (<xref ref-type="bibr" rid="ref27">27</xref>). Vitamin D from sunlight exposure also varies according to season, with a 1.7 fold higher global vitamin D deficiency rate between winter/spring compared to summer/autumn, and higher rates in people living in areas of high latitude (<xref ref-type="bibr" rid="ref1">1</xref>). An increased focus on addressing and improving vitamin D intake is needed, and the strategies and policies suggested include vitamin D supplementation, an increased intake of foods naturally high in vitamin D, traditional food fortification with vitamin D, and biofortification of vitamin D food sources (<xref ref-type="bibr" rid="ref8">8</xref>).</p>
<p>Vitamin D supplementation is important for vulnerable groups such as the elderly, and those following a diet that restricts animal foods (<xref ref-type="bibr" rid="ref1">1</xref>, <xref ref-type="bibr" rid="ref2">2</xref>, <xref ref-type="bibr" rid="ref17">17</xref>, <xref ref-type="bibr" rid="ref28">28</xref>). In Australia, vitamin D supplementation ranges from 0.6 to 17% depending on demographic group, with the highest rates among elderly women (<xref ref-type="bibr" rid="ref29">29</xref>). Low adherence within the general population and reduced accessibility by those of low socioeconomic status are significant limitations for widespread application to address vitamin D deficiency (<xref ref-type="bibr" rid="ref1">1</xref>, <xref ref-type="bibr" rid="ref2">2</xref>, <xref ref-type="bibr" rid="ref17">17</xref>, <xref ref-type="bibr" rid="ref24">24</xref>). Supplements are predominantly vitamin D<sub>3</sub>, sourced from sheep wool, which is incompatible with a vegan diet (<xref ref-type="bibr" rid="ref30">30</xref>). Vegan vitamin D<sub>3</sub> supplements from microalgae such as <italic>Nannochloropsis oceanica</italic> can be produced when they are irradiated with UV (<xref ref-type="bibr" rid="ref31">31</xref>). Similarly, mushrooms can produce vitamin D<sub>2</sub> when irradiated with UV light (<xref ref-type="bibr" rid="ref32">32</xref>) and vegan vitamin D<sub>2</sub> supplements made from mushrooms and other fungi are available. While 7-dehydrocholesterol is converted to vitamin D<sub>3</sub> in animals, ergosterol (pro-vitamin D<sub>2</sub>), which is found in mushrooms, is converted to pre-vitamin D<sub>2</sub> under UV-light, with heat required for full conversion to ergocalciferol, vitamin D<sub>2</sub> (<xref ref-type="bibr" rid="ref32">32</xref>).</p>
<p>The highest non-fortified and natural food sources of vitamin D are predominantly animal-based, such as salmon (5.4&#x2009;&#x03BC;g/100&#x2009;g) and eggs (5.9&#x2009;&#x03BC;g/100&#x2009;g) (<xref ref-type="bibr" rid="ref33">33</xref>). Animal-based solutions do not align with plant-based movements or vegan diets. A recent simulation study suggested that the achievement of adequate vitamin D intake based primarily on animal food sources is not possible within carbon emission limits (<xref ref-type="bibr" rid="ref34">34</xref>).</p>
<p>Food fortification with vitamin D has been shown to be both efficacious and cost-effective (<xref ref-type="bibr" rid="ref24">24</xref>). In Finland, voluntary vitamin D fortification of milks and margarines/fat spreads was associated with an increase in mean vitamin D status from 47.6&#x2009;nmol/L to 65.4&#x2009;nmol/L over 11&#x2009;years (<xref ref-type="bibr" rid="ref24">24</xref>). In the US, fortified milk and milk products provide the greatest contribution to dietary vitamin D intake (43.7%) (<xref ref-type="bibr" rid="ref35">35</xref>) and UV-exposed mushroom powder is approved by the FDA as a vitamin D<sub>2</sub> source for addition to some foods (<xref ref-type="bibr" rid="ref36">36</xref>). Fortified margarine is a major dietary source of vitamin D in Australia (<xref ref-type="bibr" rid="ref37">37</xref>). Australian dietary modelling showed that fortification of milk and breakfast cereals with vitamin D (1&#x2009;&#x03BC;g/100&#x2009;mL and 3.5&#x2009;&#x03BC;g/100&#x2009;g respectively) would increase average vitamin D intake from 3.6&#x2009;&#x03BC;g/day to 6.3&#x2009;&#x03BC;g/day, although remaining below IOM targets (<xref ref-type="bibr" rid="ref38">38</xref>).</p>
<p>Biofortification describes the natural vitamin D enrichment of whole foods including meats, eggs, and fish (via increased vitamin D provision to livestock), as well as UV-exposed mushrooms and yeast (<xref ref-type="bibr" rid="ref39">39</xref>, <xref ref-type="bibr" rid="ref40">40</xref>). For example, the vitamin D<sub>3</sub> content of eggs can be increased by the addition of vitamin D<sub>3</sub> to the feed of hens (<xref ref-type="bibr" rid="ref40">40</xref>). In the US, both vitamin D-enriched and sunlight-exposed mushrooms are readily available for purchase, containing 100% of the IOM EAR per 85&#x2009;g serving (<xref ref-type="bibr" rid="ref25">25</xref>, <xref ref-type="bibr" rid="ref41">41</xref>). In Australia, vitamin D-enriched mushrooms are produced via exposure to UV-lamp pulses (<xref ref-type="bibr" rid="ref42">42</xref>).</p>
<p>There is some uncertainty around the potential for excess intake of vitamin D, given the increase in nutrient availability from fortification and high-dose vitamin D supplements (<xref ref-type="bibr" rid="ref43">43</xref>), although mean intake estimates are less than 10% of the IOM upper level of intake (UL) (<xref ref-type="bibr" rid="ref1">1</xref>). An updated assessment of vitamin D intake from all sources is warranted.</p>
</sec>
<sec id="sec3">
<title>Mushrooms produce the recommended intake of vitamin D (as D<sub>2</sub>) per serve</title>
<p>The vitamin D content of UV-exposed mushrooms varies according to mushroom type, the amount of UV-exposure, the surface area exposed (whole or sliced), light intensity, and length of exposure (<xref ref-type="bibr" rid="ref28">28</xref>). Australian UV-exposed white button mushrooms can provide over 100% of vitamin D requirements in a single serve (Australian Guide to Healthy Eating, AGHE) (<xref ref-type="bibr" rid="ref28">28</xref>, <xref ref-type="bibr" rid="ref33">33</xref>). In Germany, 100&#x2009;g of sliced <italic>A. bisporus</italic> mushrooms exposed to midday, mid-summer sunlight produced 17.5&#x2009;&#x03BC;g vitamin D<sub>2</sub> after 15&#x2009;min and 32.5&#x2009;&#x03BC;g after 60&#x2009;min (<xref ref-type="bibr" rid="ref44">44</xref>), 175 and 325% of the IOM EAR, respectively (<xref ref-type="bibr" rid="ref25">25</xref>). UV-lamp pulses (1&#x2013;2&#x2009;s) after harvest produced 24&#x2009;&#x03BC;g/100&#x2009;g (240% EAR) (<xref ref-type="bibr" rid="ref33">33</xref>). The vitamin D content of UV-exposed <italic>A. bisporus</italic> mushrooms is notably higher than other dietary vitamin D sources including both oily fish (5.7&#x2009;&#x03BC;g) and eggs (7.1&#x2009;&#x03BC;g) per serve (<xref ref-type="bibr" rid="ref33">33</xref>, <xref ref-type="bibr" rid="ref45">45</xref>). Caution is warranted as there are reports of vitamin D concentrations up to 320&#x2009;&#x03BC;g/100&#x2009;g (3,200% EAR) with pulsed UV exposure, above the IOM UL (<xref ref-type="bibr" rid="ref28">28</xref>). Commercial production of vitamin D enhanced mushrooms requires standardization and testing to stay within the upper limit.</p>
<p>The Australian Food Composition Database reports that even non-UV exposed white button mushrooms can be a source of vitamin D in Australia, providing 16% of the IOM EAR per serve (<xref ref-type="bibr" rid="ref25">25</xref>, <xref ref-type="bibr" rid="ref33">33</xref>). This is in contrast to levels stated in food composition databases in the US (<xref ref-type="bibr" rid="ref46">46</xref>) and New Zealand (<xref ref-type="bibr" rid="ref47">47</xref>), at 0.02&#x2009;&#x03BC;g and 0&#x2009;&#x03BC;g vitamin D/100&#x2009;g, respectively. As mushrooms do not naturally contain vitamin D without UV exposure, it is likely that the mushrooms analyzed at point of sale in Australia had incidental UV exposure post-harvest.</p>
</sec>
<sec id="sec4">
<title>Mushrooms are a feasible and sustainable food-based source of vitamin D</title>
<p>Post synthesis, the vitamin D<sub>2</sub> content of UV-exposed <italic>A. bisporus</italic> mushrooms remains largely stable for around 7 to 10&#x2009;days when refrigerated (<xref ref-type="bibr" rid="ref28">28</xref>). The retention of vitamin D<sub>2</sub> during cooking ranges from 62 to 88%, depending on cooking method, with the highest retention in mushrooms pan-fried without oil (<xref ref-type="bibr" rid="ref48">48</xref>). UV-exposed mushrooms are therefore a feasible, food-based source of vitamin D, consumed raw or cooked. It is unknown how cooking affects the bio-accessibility of vitamin D from mushrooms in humans; this deserves further investigation as the bio-accessibility of some nutrients is enhanced following cooking.</p>
<p>UV-exposed mushrooms as a vitamin D source support sustainability efforts. The notably low environmental impact of mushrooms is largely due to their role in circular agriculture, supporting the growth, maintenance, and remediation of the surrounding environment (<xref ref-type="bibr" rid="ref49">49</xref>, <xref ref-type="bibr" rid="ref50">50</xref>). In circular agriculture, outputs from plant and animal farm waste are used as inputs in mushroom growing, and spent mushroom waste is then used to produce high-quality compost, animal feed, biofuel, and for bioremediation (<xref ref-type="bibr" rid="ref49">49</xref>, <xref ref-type="bibr" rid="ref50">50</xref>).</p>
</sec>
<sec id="sec5">
<title>Mushrooms are not considered as a vitamin D source in dietary guidelines</title>
<p>Recommended intakes for vitamin D vary worldwide; while the IOM recommends an EAR of 10&#x2009;&#x03BC;g/day, and RDA up to 20&#x2009;&#x03BC;g/day for adults over 70&#x2009;years (<xref ref-type="bibr" rid="ref25">25</xref>), Australian recommendations are based around adequate intake (AI), ranging from 5&#x2013;15&#x2009;&#x03BC;g/day depending on age group, with the highest requirements for those aged 65&#x2009;years and older (<xref ref-type="bibr" rid="ref37">37</xref>). Despite these recommendations, food-based dietary guidelines often lack provision for vitamin D; both the Australian Dietary Guidelines (ADG) and the Dietary Guidelines for Americans (DGA) (<xref ref-type="bibr" rid="ref26">26</xref>) fail to provide adequate vitamin D (<xref ref-type="bibr" rid="ref21">21</xref>, <xref ref-type="bibr" rid="ref26">26</xref>), indicating that meeting vitamin D needs is difficult and current eating patterns require additional support. Further, vitamin D does not feature as a characteristic essential nutrient in any of the ADG core food groups.</p>
<p>Similarly, the role of UV-exposed mushrooms in the provision of vitamin D is not recognized within dietary guidelines worldwide. The AGHE considers mushrooms within the &#x201C;vegetables and legumes&#x201D; subcategory of &#x201C;other vegetables,&#x201D; alongside salad vegetables such as tomatoes and cucumber (<xref ref-type="bibr" rid="ref45">45</xref>). Other vegetable subcategories include dark green or cruciferous vegetables, root vegetables, and legumes/beans (<xref ref-type="bibr" rid="ref45">45</xref>). Mushrooms are also classified as &#x201C;other vegetable&#x201D; in the DGA (<xref ref-type="bibr" rid="ref51">51</xref>). Neither the American (<xref ref-type="bibr" rid="ref52">52</xref>) nor Australian (<xref ref-type="bibr" rid="ref21">21</xref>) dietary modelling approaches that underpin guideline development considered the use of UV-exposed mushrooms as a source of vitamin D. The Australian modelling included mushrooms at around 2% of total vegetable consumption for adult diets (less than 50&#x2009;g per week), consistent with recent sales data (<xref ref-type="bibr" rid="ref53">53</xref>).</p>
</sec>
<sec id="sec6">
<title>Dietary modelling supports a role for UV-mushrooms as a key vitamin D source</title>
<p>In 2021, two dietary modelling papers from the USA (<xref ref-type="bibr" rid="ref54">54</xref>, <xref ref-type="bibr" rid="ref55">55</xref>) showed that the daily addition of an 84 gram serve of UV-exposed <italic>A. bisporus</italic> mushrooms (containing 5&#x2009;&#x03BC;g of vitamin D) dramatically improved vitamin D intakes (by 67 to 91%, depending on the baseline diet) and decreased vitamin D inadequacy in the usual US adult diet from 94.9 to 63.6%, with minimal impacts on energy and sodium levels.</p>
<p>To determine Australian-based dietary outcomes of UV-exposed mushroom addition, we modelled the effect of removing mushrooms from the &#x201C;other vegetables&#x201D; sub-category of the &#x201C;vegetables and legumes&#x201D; core food group and creating a fifth &#x201C;mushrooms only&#x201D; sub-category of vegetables and legumes. Mushrooms were then added to the diet as this fifth sub-category with increasing numbers of serves. This was carried out for three adult demographic groups (women aged 19&#x2013;30&#x2009;years, men aged 51&#x2013;70&#x2009;years, and women aged &#x003E;70&#x2009;years) and two diets (omnivore and ovo-lacto vegetarian). Methodological detail is provided in <xref ref-type="supplementary-material" rid="SM1">Supplementary Figure S1</xref>.</p>
<p>The modelling showed notable increases in vitamin D intakes, as well as several additional micronutrients, for all demographic groups and both diet models (omnivore and ovo-lacto vegetarian). While all baseline diets were inadequate for vitamin D, ranging from 10 to 31% of the IOM RDA across demographic groups, the addition of one serve/day (75&#x2009;g) of UV-exposed mushrooms enabled all demographic groups to exceed their recommended dietary vitamin D intake by 28 to 87% (<xref ref-type="fig" rid="fig1">Figure 1</xref>). Recommendations were achieved at a minimum of 4 serves per week for adults up to the age of 70&#x2009;years; over this age, 5 and 6 serves per week for the omnivore and ovo-lacto vegetarian diet, respectively, were required (data not shown). Beneficial effects on additional micronutrients included an 18.6 to 34.2% increase in selenium across all demographic groups and both diets (<xref ref-type="supplementary-material" rid="SM1">Supplementary Table S1</xref>). While there were small decreases in some nutrients in substitution models, such as riboflavin and vitamin B12, NRVs were still met. There was a negligible impact on energy intakes in all diets, even when 7 serves of UV-exposed mushrooms were added to current recommendations per week.</p>
<fig position="float" id="fig1">
<label>Figure 1</label>
<caption>
<p>Vitamin D intakes as a percentage of the Institute of Medicine Recommended Daily Allowance for vitamin D for each demographic group, as a result of dietary modelling focused on mushrooms as a separate subcategory of vegetables within Foundation Diets from the Australian Dietary Guidelines. One serve is equivalent to 75&#x2009;g.</p>
</caption>
<graphic xlink:href="fnut-11-1384273-g001.tif"/>
</fig>
<p>The findings show that UV-exposed mushrooms have the potential to make a meaningful contribution to vitamin D intakes of Australian adults, allowing an individual to meet their vitamin D needs if consumed daily. However, the modelled intake (525&#x2009;g/week) is notably higher than current intakes, both in Australia (50&#x2009;g/week) (<xref ref-type="bibr" rid="ref53">53</xref>) and globally (100&#x2009;g/week) (<xref ref-type="bibr" rid="ref56">56</xref>). Education, policy, and programs on the benefits of UV-exposed mushrooms alongside their production across major suppliers may offer an effective solution to addressing low vitamin D intakes and inadequacy. Although the consumption of UV-exposed mushrooms may not, in practice, consistently align with the amounts necessary to fully provide for recommended vitamin D intakes, UV-exposed mushrooms can play an important role as part of the solution to low vitamin D intakes, supporting other dietary sources of vitamin D, as well as supplementation where necessary. This is particularly important for those consuming vegetarian and vegan diets, where there is a low intake of animal-based vitamin D. Future dietary modelling in dietary guidelines needs to consider UV-exposed mushrooms as a source of vitamin D.</p>
<p>The modelling approach presented here was focused on adult demographic groups only; however, UV-exposed mushrooms may also represent an additional vitamin D source for children. While mushrooms contain insoluble fibres such as chitin, and excessive intakes of fibre may be associated with gastrointestinal discomfort in this population (<xref ref-type="bibr" rid="ref57">57</xref>), the prevalence of vitamin D deficiency in Australian children is lower than that in adults (<xref ref-type="bibr" rid="ref58">58</xref>), indicating that a reduced consumption of UV-exposed mushrooms may support adequate intakes. Further research to determine consumption levels in children and effect on vitamin D status is warranted.</p>
</sec>
<sec id="sec7">
<title>Is D<sub>2</sub> from mushrooms a substitute for D<sub>3</sub>?</title>
<p>The efficacy of vitamin D<sub>2</sub> compared to D<sub>3</sub> for increasing vitamin D status (25-hydroxyvitamin D, 25 (OH)D) is yet to be fully understood. While meta-analyses have shown that vitamin D<sub>3</sub> is more effective than vitamin D<sub>2</sub> in increasing total vitamin D status, these relationships appear to be modified by both BMI and baseline vitamin D status (<xref ref-type="bibr" rid="ref59">59</xref>). A 2023 systematic literature review identified that vitamin D<sub>2</sub> from UV-exposed mushrooms (from 8.8&#x2009;&#x03BC;g/day) consistently increased serum levels of vitamin D<sub>2</sub> compared to placebo (<xref ref-type="bibr" rid="ref60">60</xref>). However, there was no change in total vitamin D levels in most studies, possibly explained by the concomitant decrease in vitamin D<sub>3</sub> levels in 50% of studies. This may reflect a tight regulation of total vitamin D levels (<xref ref-type="bibr" rid="ref61">61</xref>). In one trial, there was a decrease in total vitamin D that was greater in subjects with higher vitamin D at baseline (<xref ref-type="bibr" rid="ref62">62</xref>). A parallel RCT not included in the 2023 review found that consumption of mushrooms containing D<sub>2</sub> was as effective at increasing and maintaining total serum vitamin D levels as both supplemental vitamin D<sub>2</sub> and D<sub>3</sub> (all 50&#x2009;&#x03BC;g/day) (<xref ref-type="bibr" rid="ref63">63</xref>). In this study, baseline vitamin D was bordering deficiency. A 2024 systematic review with meta-analysis found no effect of mushroom vitamin D<sub>2</sub> on serum vitamin D status, although significance was borderline (<italic>p</italic>&#x2009;=&#x2009;0.06) and a statistically and clinically significant increase was seen in sub-analyses of the lowest (42&#x2009;nmol/L) vs. highest (&#x003E;75&#x2009;nmol/L) baseline vitamin D status (<xref ref-type="bibr" rid="ref64">64</xref>). Similarly, vitamin D<sub>2</sub> supplementation-induced decreases in vitamin D<sub>3</sub> appear to be highest in those with highest baseline vitamin D (<xref ref-type="bibr" rid="ref64">64</xref>). The relative effectiveness of vitamin D<sub>2</sub> and D<sub>3</sub> for increasing vitamin D status also appears to depend on level and frequency of dose, with larger differences between D<sub>2</sub> and D<sub>3</sub> in bolus compared to daily dosing protocols (<xref ref-type="bibr" rid="ref65">65</xref>, <xref ref-type="bibr" rid="ref66">66</xref>). Despite these differences, pharmacologic doses of vitamin D<sub>2</sub> have shown the ability to maintain serum vitamin D above 50&#x2009;nmol/L in clinical settings of vitamin D deficiency (<xref ref-type="bibr" rid="ref67">67</xref>, <xref ref-type="bibr" rid="ref68">68</xref>). Together, the evidence suggests that beneficial increases in vitamin D status can occur with vitamin D<sub>2</sub> from mushrooms in those with deficient or insufficient vitamin D status.</p>
<p>A key aspect of vitamin D biology that requires further understanding is the physiological role of D<sub>2</sub> vs. D<sub>3</sub>. While sharing a similar structure, differences in the half-life of the hydroxylated forms of vitamin D<sub>2</sub> and D<sub>3</sub> have been identified (D<sub>2</sub> is shorter), at least in some populations (<xref ref-type="bibr" rid="ref69">69</xref>). There may also be differences in the rate of hydroxylation, affinity for the vitamin D binding protein, and binding to the vitamin D receptor (<xref ref-type="bibr" rid="ref61">61</xref>). A recent analysis of the blood transcriptome following D<sub>2</sub> vs. D<sub>3</sub> supplementation showed that, despite there being overlap in gene expression changes, some were specific to one form of the vitamin vs. the other (<xref ref-type="bibr" rid="ref70">70</xref>).</p>
<p>Together, the findings suggest that while vitamin D<sub>2</sub> is not a direct substitute for vitamin D<sub>3</sub>, vitamin D enhanced mushrooms can increase vitamin D status among those who are deficient or insufficient, those with low potential for UV exposure, and those with limited intake of animal sources of vitamin D. A more comprehensive analysis of the biological effects of the two forms of vitamin D in humans is needed.</p>
</sec>
<sec id="sec8">
<title>Call to action: promoting UV-exposed mushrooms as part of the solution to vitamin D deficiency</title>
<p>UV-exposed mushrooms show potential to be a meaningful, whole-food, and vegan source of vitamin D. Current dietary guidelines are increasingly focused on environmental sustainability and &#x201C;plant-based&#x201D; diets; while possessing numerous benefits, such guidelines may inadvertently increase vitamin D deficiency and suboptimal intakes of other micronutrients, such as selenium.</p>
<p>There is an opportunity for health professionals, stakeholders, and policy makers to provide greater guidance on maximizing diet as a source of vitamin D, by UV-exposing mushrooms and increasing their focus in dietary guidelines worldwide. Moving mushrooms into a new sub-category of the vegetables core food group, as we have modelled, could facilitate an increase in mushroom intake. Practical guidance in using and cooking mushrooms as a major source of vitamin D can be provided, such as purchasing UV-exposed mushrooms or putting mushrooms in the sun prior to eating (15&#x2009;min in the sun between 10&#x2009;am and 3&#x2009;pm, and store in fridge for up to 8&#x2009;days (<xref ref-type="bibr" rid="ref28">28</xref>)).</p>
<p>The production of UV-exposed mushrooms is limited and comes at a cost to farmers and consumers, creating a barrier for its inclusion. Government action in supporting the universal UV-exposure of mushrooms, similarly to efforts in the universal iodization of salt (<xref ref-type="bibr" rid="ref71">71</xref>), could enhance efforts to address vitamin D deficiency globally. UV-exposed mushrooms can contribute to addressing vitamin D inadequacies in a sustainable, whole-food fashion, warranting their consideration as a key and substantial dietary source of vitamin D.</p>
</sec>
</sec>
<sec sec-type="data-availability" id="sec9">
<title>Data availability statement</title>
<p>The original contributions presented in the study are included in the article/<xref ref-type="supplementary-material" rid="SM1">Supplementary material</xref>, further inquiries can be directed to the corresponding author.</p>
</sec>
<sec sec-type="ethics-statement" id="sec10">
<title>Ethics statement</title>
<p>Ethical approval was not required for the study involving humans in accordance with the local legislation and institutional requirements. Written informed consent to participate in this study was not required from the participants or the participants&#x2019; legal guardians/next of kin in accordance with the national legislation and the institutional requirements.</p>
</sec>
<sec sec-type="author-contributions" id="sec11">
<title>Author contributions</title>
<p>CS: Conceptualization, Data curation, Formal analysis, Investigation, Methodology, Writing &#x2013; original draft, Writing &#x2013; review &#x0026; editing. TC: Conceptualization, Methodology, Writing &#x2013; review &#x0026; editing. JW: Writing &#x2013; review &#x0026; editing. PP: Methodology, Validation, Writing &#x2013; review &#x0026; editing. EB: Writing &#x2013; review &#x0026; editing. FF-M: Funding acquisition, Methodology, Supervision, Writing &#x2013; review &#x0026; editing.</p>
</sec>
</body>
<back>
<sec sec-type="funding-information" id="sec12">
<title>Funding</title>
<p>The author(s) declare that financial support was received for the research, authorship, and/or publication of this article. This project has been funded by a research grant from Horticulture Innovation Australia. The funding body had no contribution to the modelling methodology, the data analysis, drafting of the manuscript, nor interpretation of findings.</p>
</sec>
<ack>
<p>The authors would like to acknowledge Leah Bramich, Australian Mushroom Growers&#x2019; Association, and Jacqui Simpson from Horticulture Innovation Australia, who provided top-line suggestions on content.</p>
</ack>
<sec sec-type="COI-statement" id="sec13">
<title>Conflict of interest</title>
<p>CS, TC, JW, EB, and FF-M were employed by FOODiQ Global.</p>
<p>The remaining author declares that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p>
</sec>
<sec id="sec100" sec-type="disclaimer">
<title>Publisher&#x2019;s note</title>
<p>All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.</p>
</sec>
<sec sec-type="supplementary-material" id="sec14">
<title>Supplementary material</title>
<p>The Supplementary material for this article can be found online at: <ext-link xlink:href="https://www.frontiersin.org/articles/10.3389/fnut.2024.1384273/full#supplementary-material" ext-link-type="uri">https://www.frontiersin.org/articles/10.3389/fnut.2024.1384273/full#supplementary-material</ext-link></p>
<supplementary-material xlink:href="Data_Sheet_1.pdf" id="SM1" mimetype="application/pdf" xmlns:xlink="http://www.w3.org/1999/xlink"/>
</sec>
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