At the end of December 2015, Apple Inc. (NASDAQ: AAPL) reported 5.54 billion shares of stock issued and outstanding. In the company’s proxy statement filed in January, Apple listed just two firms holding more than 5% of its outstanding shares.

To meet that requirement, a shareholder would need to own 277.22 million shares. At the December 28, 2015, date Apple used to calculate outstanding shares, the stock closed at $106.82, indicating that a 5% investment was then worth about $29.6 billion. At Wednesday’s closing price of $105.79, a 5% stake in Apple was worth $29.3 billion.

Now that Apple stock is approaching its end-of-2015 level, it’s interesting to note that the stock was the most heavily purchased stock in the fourth calendar quarter of 2015 and the most heavily sold in the first quarter of 2016. The top 50 hedge funds sold some $7.1 billion worth of Apple stock in the first quarter, compared with purchases of $2.2 billion in the fourth quarter of 2015.

A $4.8 billion sale by activist investor Carl Icahn was responsible for most of the first-quarter selling of Apple stock, and those who held on until mid-April saw a gain of about 6% in the first three and a half months of this year. So who were the lucky souls?

Here are Apple’s top five institutional shareholders:

At the end of December 2015, Vanguard Group beneficially owned 332.24 million shares (5.99%) of Apple stock, valued at about $35.15 billion at last night’s closing price. The investment firm also holds large stakes in Microsoft ($27.4 billion), Exxon Mobil ($22.35 billion) and Johnson & Johnson ($19.54 billion).

BlackRock and several affiliates owned 5.7% of Apple’s outstanding shares (315,862,269), valued at around $33.41 billion. According to Holdings Channel, BlackRock Institutional Trust Company owned nearly half the total and Apple remains the trust’s largest position based on value. The trust also holds large positions in Microsoft ($11.5 billion), Exxon Mobil ($9.29 billion) and Johnson & Johnson ($7.96 billion).

State Street owned 146.61 million shares of Apple stock as of March 31, 2016. That 2.64% stake was valued at around $22.81 billion at Wednesday’s closing price. Apple is State Street’s largest holding, followed by Microsoft ($16.96 billion), Johnson & Johnson ($15.7 billion) and Exxon Mobil ($15.6 billion).

FMR, a Fidelity company, owned 2.62% (145.17 million) of Apple’s outstanding shares. At Wednesday’s closing price, its stake is valued at $15.36 billion. FMR’s largest holdings include nearly $26 billion stakes in both classes of Google stock, $17.4 billion in Facebook stock and $11.75 billion in Amazon stock.

Northern Trust owned 70.3 million shares of Apple stock (1.26%), valued at $7.44 billion. The firm’s other large holdings include Microsoft ($5.3 billion), Exxon Mobil ($4.49 billion) and GE ($3.85 billion).

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