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

    <schemaVersion>X0206</schemaVersion>

    <documentType>3</documentType>

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

    <noSecuritiesOwned>0</noSecuritiesOwned>

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

    <reportingOwner>
        <reportingOwnerId>
            <rptOwnerCik>0001543112</rptOwnerCik>
            <rptOwnerName>Morningside Venture Investments Ltd</rptOwnerName>
        </reportingOwnerId>
        <reportingOwnerAddress>
            <rptOwnerStreet1>C/O THC MANAGEMENT SERVICES S.A.M.</rptOwnerStreet1>
            <rptOwnerStreet2>3-5 AVENUE DES CITRONNIERS</rptOwnerStreet2>
            <rptOwnerCity>MONACO</rptOwnerCity>
            <rptOwnerState>O9</rptOwnerState>
            <rptOwnerZipCode>MC 98000</rptOwnerZipCode>
            <rptOwnerStateDescription>MONACO</rptOwnerStateDescription>
        </reportingOwnerAddress>
        <reportingOwnerRelationship>
            <isDirector>0</isDirector>
            <isOfficer>0</isOfficer>
            <isTenPercentOwner>1</isTenPercentOwner>
            <isOther>0</isOther>
        </reportingOwnerRelationship>
    </reportingOwner>

    <reportingOwner>
        <reportingOwnerId>
            <rptOwnerCik>0001914583</rptOwnerCik>
            <rptOwnerName>MVIL, LLC</rptOwnerName>
        </reportingOwnerId>
        <reportingOwnerAddress>
            <rptOwnerStreet1>C/O 22ND FLOOR, HANG LUNG CENTRE</rptOwnerStreet1>
            <rptOwnerStreet2>2-20 PATERSON ST.</rptOwnerStreet2>
            <rptOwnerCity>CAUSEWAY BAY, HONG KONG</rptOwnerCity>
            <rptOwnerState>K3</rptOwnerState>
            <rptOwnerZipCode></rptOwnerZipCode>
            <rptOwnerStateDescription>HONG KONG</rptOwnerStateDescription>
        </reportingOwnerAddress>
        <reportingOwnerRelationship>
            <isDirector>0</isDirector>
            <isOfficer>0</isOfficer>
            <isTenPercentOwner>1</isTenPercentOwner>
            <isOther>0</isOther>
        </reportingOwnerRelationship>
    </reportingOwner>

    <derivativeTable>
        <derivativeHolding>
            <securityTitle>
                <value>Class A Common Stock</value>
            </securityTitle>
            <conversionOrExercisePrice>
                <footnoteId id="F1"/>
            </conversionOrExercisePrice>
            <exerciseDate>
                <footnoteId id="F1"/>
            </exerciseDate>
            <expirationDate>
                <footnoteId id="F1"/>
            </expirationDate>
            <underlyingSecurity>
                <underlyingSecurityTitle>
                    <value>Common Stock</value>
                </underlyingSecurityTitle>
                <underlyingSecurityShares>
                    <value>12857931</value>
                </underlyingSecurityShares>
            </underlyingSecurity>
            <ownershipNature>
                <directOrIndirectOwnership>
                    <value>D</value>
                    <footnoteId id="F2"/>
                </directOrIndirectOwnership>
            </ownershipNature>
        </derivativeHolding>
        <derivativeHolding>
            <securityTitle>
                <value>Class A Common Stock</value>
            </securityTitle>
            <conversionOrExercisePrice>
                <footnoteId id="F1"/>
            </conversionOrExercisePrice>
            <exerciseDate>
                <footnoteId id="F1"/>
            </exerciseDate>
            <expirationDate>
                <footnoteId id="F1"/>
            </expirationDate>
            <underlyingSecurity>
                <underlyingSecurityTitle>
                    <value>Common Stock</value>
                </underlyingSecurityTitle>
                <underlyingSecurityShares>
                    <value>1000000</value>
                </underlyingSecurityShares>
            </underlyingSecurity>
            <ownershipNature>
                <directOrIndirectOwnership>
                    <value>D</value>
                    <footnoteId id="F2"/>
                    <footnoteId id="F3"/>
                </directOrIndirectOwnership>
            </ownershipNature>
        </derivativeHolding>
    </derivativeTable>

    <footnotes>
        <footnote id="F1">The shares of Class A common stock automatically convert into the same number of shares of the Issuer's Common Stock without payment or further consideration in connection with the closing of the Issuer's business combination transaction.  The Class A Common Stock has no expiration date.</footnote>
        <footnote id="F2">Frances Anne Elizabeth Richard, Jill Marie Franklin, Peter Stuart Allenby Edwards and Cheung Ka Ho are the directors of Morningside and share voting and dispositive power with respect to the securities held by Morningside Venture Investments Limited (&quot;Morningside&quot;) and MVIL, LLC (&quot;MVIL LLC&quot;), Morningside's wholly-owned subsidiary.  Ms. Richard, Ms. Franklin, Mr. Edwards and Mr. Cheung each disclaim ownership of the securities owned by Morningside and MVIL LLC.  Cheng Yee Wing Betty and Wong See Wai are the managers of MVIL and share voting and dispositive power with respect to the securities held by MVIL LLC.  Ms. Cheng and Mr. Wong each disclaim ownership of the securities owned by MVIL LLC.</footnote>
        <footnote id="F3">Represent shares held by MVIL, LLC.</footnote>
    </footnotes>

    <remarks>The Reporting Persons are jointly filing this Form 3 pursuant to Rule 16a-3(j) under the Securities Exchange Act of 1934.</remarks>

    <ownerSignature>
        <signatureName>/s/ Frances Anne Elizabeth Richards, for Morningside Venture Investments Limited</signatureName>
        <signatureDate>2022-03-17</signatureDate>
    </ownerSignature>

    <ownerSignature>
        <signatureName>/s/ Chen Yee Wing Betty, for MVIL LLC</signatureName>
        <signatureDate>2022-03-17</signatureDate>
    </ownerSignature>
</ownershipDocument>
