<?xml version="1.0"?>
<ownershipDocument>

    <schemaVersion>X0306</schemaVersion>

    <documentType>4</documentType>

    <periodOfReport>2022-02-02</periodOfReport>

    <issuer>
        <issuerCik>0001822691</issuerCik>
        <issuerName>GreenLight Biosciences Holdings, PBC</issuerName>
        <issuerTradingSymbol>GRNA</issuerTradingSymbol>
    </issuer>

    <reportingOwner>
        <reportingOwnerId>
            <rptOwnerCik>0001908721</rptOwnerCik>
            <rptOwnerName>Walker Matthew Alan</rptOwnerName>
        </reportingOwnerId>
        <reportingOwnerAddress>
            <rptOwnerStreet1>C/O GREENLIGHT BIOSCIENCES HOLDINGS, PBC</rptOwnerStreet1>
            <rptOwnerStreet2>200 BOSTON AVENUE, SUITE 3100</rptOwnerStreet2>
            <rptOwnerCity>MEDFORD</rptOwnerCity>
            <rptOwnerState>MA</rptOwnerState>
            <rptOwnerZipCode>02155</rptOwnerZipCode>
            <rptOwnerStateDescription></rptOwnerStateDescription>
        </reportingOwnerAddress>
        <reportingOwnerRelationship>
            <isDirector>1</isDirector>
            <isOfficer>0</isOfficer>
            <isTenPercentOwner>1</isTenPercentOwner>
            <isOther>0</isOther>
        </reportingOwnerRelationship>
    </reportingOwner>

    <reportingOwner>
        <reportingOwnerId>
            <rptOwnerCik>0001785504</rptOwnerCik>
            <rptOwnerName>BUILDERS VISION LLC</rptOwnerName>
        </reportingOwnerId>
        <reportingOwnerAddress>
            <rptOwnerStreet1>110 N.W. 2ND STREET</rptOwnerStreet1>
            <rptOwnerStreet2>SUITE 300</rptOwnerStreet2>
            <rptOwnerCity>BENTONVILLE</rptOwnerCity>
            <rptOwnerState>AR</rptOwnerState>
            <rptOwnerZipCode>72172</rptOwnerZipCode>
            <rptOwnerStateDescription></rptOwnerStateDescription>
        </reportingOwnerAddress>
        <reportingOwnerRelationship>
            <isDirector>0</isDirector>
            <isOfficer>0</isOfficer>
            <isTenPercentOwner>1</isTenPercentOwner>
            <isOther>0</isOther>
        </reportingOwnerRelationship>
    </reportingOwner>

    <nonDerivativeTable>
        <nonDerivativeTransaction>
            <securityTitle>
                <value>Common Stock</value>
            </securityTitle>
            <transactionDate>
                <value>2022-02-02</value>
            </transactionDate>
            <transactionCoding>
                <transactionFormType>4</transactionFormType>
                <transactionCode>A</transactionCode>
                <equitySwapInvolved>0</equitySwapInvolved>
            </transactionCoding>
            <transactionAmounts>
                <transactionShares>
                    <value>14343021</value>
                </transactionShares>
                <transactionPricePerShare>
                    <footnoteId id="F1"/>
                </transactionPricePerShare>
                <transactionAcquiredDisposedCode>
                    <value>A</value>
                </transactionAcquiredDisposedCode>
            </transactionAmounts>
            <postTransactionAmounts>
                <sharesOwnedFollowingTransaction>
                    <value>14343021</value>
                </sharesOwnedFollowingTransaction>
            </postTransactionAmounts>
            <ownershipNature>
                <directOrIndirectOwnership>
                    <value>I</value>
                </directOrIndirectOwnership>
                <natureOfOwnership>
                    <value>by Builders Vision, LLC</value>
                    <footnoteId id="F2"/>
                    <footnoteId id="F3"/>
                </natureOfOwnership>
            </ownershipNature>
        </nonDerivativeTransaction>
        <nonDerivativeTransaction>
            <securityTitle>
                <value>Common Stock</value>
            </securityTitle>
            <transactionDate>
                <value>2022-02-02</value>
            </transactionDate>
            <transactionCoding>
                <transactionFormType>4</transactionFormType>
                <transactionCode>P</transactionCode>
                <equitySwapInvolved>0</equitySwapInvolved>
            </transactionCoding>
            <transactionAmounts>
                <transactionShares>
                    <value>1500000</value>
                </transactionShares>
                <transactionPricePerShare>
                    <value>10</value>
                    <footnoteId id="F4"/>
                </transactionPricePerShare>
                <transactionAcquiredDisposedCode>
                    <value>A</value>
                </transactionAcquiredDisposedCode>
            </transactionAmounts>
            <postTransactionAmounts>
                <sharesOwnedFollowingTransaction>
                    <value>15843021</value>
                </sharesOwnedFollowingTransaction>
            </postTransactionAmounts>
            <ownershipNature>
                <directOrIndirectOwnership>
                    <value>I</value>
                </directOrIndirectOwnership>
                <natureOfOwnership>
                    <value>by Builders Vision, LLC</value>
                    <footnoteId id="F2"/>
                    <footnoteId id="F3"/>
                </natureOfOwnership>
            </ownershipNature>
        </nonDerivativeTransaction>
    </nonDerivativeTable>

    <footnotes>
        <footnote id="F1">Issued pursuant to the terms of a business combination agreement involving the issuer and GreenLight Biosciences, Inc. in exchange for capital stock previously issued by such company.</footnote>
        <footnote id="F2">Shares owned directly: 3,673,694 by S2G Builders Food &amp; Agriculture Fund III, L.P. (&quot;Fund III&quot;); 2,087,043 by S2G Ventures Fund I, L.P. (&quot;Fund I&quot;); and 8,582,284 by S2G Ventures Fund II, L.P. (&quot;Fund II&quot;). Builders Vision, LLC is Manager of Funds I and II, and General Partner of Fund III, and has power to vote or direct the voting of shares held by the funds. General Partners of Fund I and Fund II are S2G Ventures, LLC and S2G Ventures II, LLC, respectively.</footnote>
        <footnote id="F3">(Continued) Mr. Walker, a director of the Issuer, is a Managing Director at Builders Vision, LLC, the impact platform founded by Lukas T. Walton. By virtue of the foregoing, S2G Ventures LLC, S2G Ventures II, LLC, and Mr. Walton may be deemed to indirectly beneficially own (as defined in Rule 13d-3 of the Exchange Act) the shares of common stock held by the respective funds. Mr. Walker and Mr. Walton each disclaims beneficial ownership of these shares of common stock except to the extent of any pecuniary interest therein.</footnote>
        <footnote id="F4">Amount consists of 1,500,000 shares of common stock issued pursuant to a subscription agreement entered into between the Issuer and S2G Builders Food &amp; Agriculture Fund III, L.P.</footnote>
    </footnotes>

    <remarks>S2G Advisors, LLC has changed its name to Builders Vision, LLC.

Mr. Walker's middle name is spelled Allen, not Alan.</remarks>

    <ownerSignature>
        <signatureName>/s/ David Kennedy, Attorney-in-fact for Matthew Allen Walker</signatureName>
        <signatureDate>2022-02-04</signatureDate>
    </ownerSignature>

    <ownerSignature>
        <signatureName>/s/ Lisa J. Forbes, as authorized officer of Builders Vision, LLC</signatureName>
        <signatureDate>2022-02-04</signatureDate>
    </ownerSignature>
</ownershipDocument>
