Case: 19-2255    Document: 58     Page: 1   Filed: 08/03/2020




   United States Court of Appeals
       for the Federal Circuit
                  ______________________

        BIO-RAD LABORATORIES, INC., THE
            UNIVERSITY OF CHICAGO,
                Plaintiffs-Appellees

                             v.

                  10X GENOMICS INC.,
                   Defendant-Appellant
                  ______________________

                   2019-2255, 2019-2285
                  ______________________

     Appeals from the United States District Court for the
 District of Delaware in No. 1:15-cv-00152-RGA, Judge
 Richard G. Andrews.
                 ______________________

                 Decided: August 3, 2020
                 ______________________

     EDWARD R. REINES, Weil, Gotshal & Manges LLP, Red-
 wood Shores, CA, argued for plaintiffs-appellees. Also rep-
 resented by CHRISTOPHER SHAWN LAVIN, DEREK C.
 WALTER.

     E. JOSHUA ROSENKRANZ, Orrick, Herrington & Sutcliffe
 LLP, New York, NY, argued for defendant-appellant. Also
 represented by ELIZABETH MOULTON, Menlo Park, CA;
 MELANIE L. BOSTWICK, Washington, DC; AZRA
 HADZIMEHMEDOVIC, Tensegrity Law Group LLP, McLean,
 VA; MATTHEW D. POWERS, ROBERT LEWIS GERRITY,
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 2           BIO-RAD LABORATORIES, INC.   v. 10X GENOMICS INC.



 Redwood Shores, CA.

    STEVEN R. TRYBUS, Locke Lord LLP, Chicago, IL, for
 amicus curiae The Broad Institute, Inc.
                 ______________________

 Before NEWMAN, O’MALLEY, and TARANTO, Circuit Judges.
 O’MALLEY, Circuit Judge.
      Bio-Rad Laboratories, Inc. and the University of Chi-
 cago (collectively, “Bio-Rad”), accused 10X Genomics Inc.
 (“10X”) of infringing three patents: U.S. Patent Nos.
 8,889,083 (“’083 patent”); 8,304,193 (“’193 patent”); and
 8,329,407 (“’407 patent”). The United States District Court
 for the District of Delaware held a jury trial in November
 2018. The jury found all three patents valid and willfully
 infringed. It also awarded damages in the amount of
 $23,930,716. Post-trial, the district court denied 10X’s mo-
 tion for judgment as a matter of law (“JMOL”) under Fed-
 eral Rule of Civil Procedure 50(b), rejecting 10X’s
 arguments that (1) the accused products do not infringe; (2)
 10X’s infringement was not willful; (3) the asserted claims
 are invalid; and (4) Bio-Rad had failed to present a legally
 sufficient damages case. Bio-Rad Labs. Inc. v. 10X Ge-
 nomics, Inc., 396 F. Supp. 3d 369 (D. Del. 2019). The dis-
 trict court also granted Bio-Rad’s motion for a permanent
 injunction. Bio-Rad Labs. Inc. v. 10X Genomics, Inc., No.
 15-cv-152-RGA, 2019 WL 3322322, at *1 (D. Del. July 24,
 2019). This appeal followed. For the reasons discussed be-
 low, we affirm-in-part, reverse-in-part, vacate-in-part, and
 remand. Specifically, we affirm the judgment of infringe-
 ment of the ’083 patent and the entirety of the jury’s dam-
 ages award. We reverse, however, the district court’s
 construction of the asserted claims of the ’407 and ’193 pa-
 tents and vacate the judgment of infringement of those pa-
 tents. We remand for a new trial on the issue of whether
 10X’s accused products infringe the ’407 and ’193 patents
 under the proper claim construction. We also vacate the
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 district court’s injunction, but only with respect to 10X’s
 Linked-Reads and CNV product lines.
                       I. BACKGROUND
                A. The Claimed Technology and
                      the Patents-in-Suit
      The patents-in-suit are directed to systems and meth-
 ods for forming microscopic droplets (also called “plugs”) of
 fluids to perform biochemical reactions. Microfluidic sys-
 tems—often called “labs-on-a-chip”—allow scientists to
 conduct microscale chemical and biological reactions. For
 example, the technology allows scientists to analyze and
 compare DNA, RNA, and proteins within large numbers of
 individual cells. This technology therefore has applica-
 tions in medical diagnostics and high-throughput screen-
 ing.
     Microfluidic systems utilize chips that have “microflu-
 idic channels,” hair-width pathways through which cells
 and fluids flow. In these systems, biological samples can
 be partitioned into single-cell-width droplets, which func-
 tion as mini-test tubes. Each droplet holds a single cell and
 the required reagents for the biochemical reaction. Drop-
 lets are formed by “pinching off”—flowing a carrier-fluid
 and substrate/plug-fluid (which are immiscible with each
 other) through the microfluid channels and applying pres-
 sure. The biochemical reactions may occur “on chip,” i.e.,
 in the channels inside the microchips, or, the droplets may
 be collected to allow the reactions to occur “off chip.”
     The parties agree that claim 1 of the ’083 patent, copied
 below, is representative.
    1. A microfluidic system comprising:
    a non-fluorinated microchannel;
    a carrier fluid comprising a fluorinated oil and a
    fluorinated surfactant comprising a hydrophilic
    head group in the microchannel;
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     at least one plug comprising an aqueous plug-fluid
     in the microchannel and substantially encased by
     the carrier-fluid, wherein the fluorinated surfactant
     is present at a concentration such that surface ten-
     sion at the plug-fluid/microchannel wall interface is
     higher than surface tension at the plug-fluid/carrier
     fluid interface.
 ’083 patent, claim 1.
     During prosecution of the ’083 patent, the inventors
 amended the claims to overcome a rejection based on the
 prior art U.S. Patent No. 7,294,503 (“Quake”). Quake dis-
 closed microchannels formed or coated with Teflon (a fluor-
 inated polymer) or other fluorinated oils. The inventors
 distinguished the prior art by arguing that, unlike Quake,
 the as-filed application for the ’083 patent attempts to pre-
 vent droplets from sticking to the walls of microchannels
 and requires that the “surfactant should be chemically sim-
 ilar to the carrier fluid and chemically different from the
 channel walls.” J.A. 16640. The inventors amended the
 claims to require non-fluorinated microchannels and a
 fluorinated surfactant, which would not react with each
 other. 1 They explained that, as amended, the claims were


     1   The amendment at issue added the claim limita-
 tions shown in underlined text below:
     A microfluidic system comprising:
     a non-fluorinated microchannel;
     a carrier fluid comprising a fluorinated oil and a
     fluorinated surfactant comprising a hydrophilic
     head group in the microchannel;
     at least one plug comprising an aqueous plug-fluid
     in the microchannel and substantially encased by
     the carrier-fluid, wherein the fluorinated surfactant
     is present at a concentration such that surface
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 distinct from Quake, which did not teach microchannels
 and carrier fluids that were chemically distinct. Rather, in
 their view, Quake taught coating the microchannels with a
 fluorinated oil and using fluorinated surfactants in the car-
 rier fluid. The fluorinated microchannels and surfactants
 could, therefore, react with each other.
     The other two asserted patents, the ’407 and the ’193
 patents, are continuations of the same parent application.
 Claim 1 of the ’407 patent is reproduced below:
    1. A method for conducting a reaction in plugs in a
    microfluidic system, comprising the steps of:
    providing the microfluidic system comprising at
    least two channels having at least one junction;
    continuously flowing an aqueous fluid containing at
    least one biological molecule and at least one rea-
    gent for conducting the reaction between the biolog-
    ical molecule and the at least one reagent through
    a first channel of the at least two channels;
    continuously flowing a carrier fluid immiscible with
    the aqueous fluid through the second channel of the
    at least two channels;
    forming at least one plug of the aqueous fluid con-
    taining the at least one biological molecule and the
    at least one reagent by partitioning the aqueous
    fluid with the flowing immiscible carrier fluid at the
    junction of the at least two channels, the plug being
    substantially surrounded by the immiscible carrier
    fluid flowing through the channel, wherein the at



    tension at the plug-fluid/microchannel wall inter-
    face is higher than surface tension at the plug-
    fluid/carrier fluid interface.
 J.A. 16635.
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     least one plug comprises at least one biological mol-
     ecule and the at least one reagent for conducting the
     reaction with the at least one biological molecule;
     and
     providing conditions suitable for the reaction in the
     at least one plug involving the at least one biological
     molecule and the at least one reagent to form a re-
     action product.
 ’407 patent, claim 1.
     The only independent claim of the ’193 patent is iden-
 tical to claim 1 of the ’407 patent, except that it specifies
 “an autocatalytic reaction” instead of a biological reaction.
 10X maintains that its arguments regarding the ’407 pa-
 tent “apply equally to the ’193 patent unless otherwise
 noted.” Appellant’s Br. 44 n.3.
                   B. The Accused Products
      10X has five accused product lines: Single Cell 3’ Gene
 Expression, Linked-Reads, Single Cell V(D)J, Single Cell
 ATAC-seq, and Single Cell CNV. 10X claims to have suc-
 cessfully invented non-infringing alternatives for three of
 its five product lines but has not yet been able to design a
 replacement for two: Linked-Reads and Single Cell CNV.
     Each product line uses a hardware instrument, micro-
 fluidic chips, and a variety of specialized reagents. The
 hardware instrument is called a “controller.” The disposa-
 ble microfluidic chips, which fit in the instrument, have
 networks of “microfluidic channels,” each about the width
 of a human hair. In the accused products, droplets are
 formed at junctions in the microfluidic channels. The rea-
 gents encompass a variety of products such as enzymes,
 DNA barcodes, and 10X’s proprietary microscopic beads.
     The record does not establish which 10X products cor-
 respond to each of the product lines. For the ’083 patent,
 there were 6 accused products: Chromium Genome/Exome;
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 BIO-RAD LABORATORIES, INC.   v. 10X GENOMICS INC.           7



 Chromium Genome/Exome with Kynar; GemCode Long
 Read; Chromium Single Cell 3’; Chromium Single Cell 3’
 with Kynar; and Chromium Single Cell V(D)J with Kynar.
 The jury found that all six products infringe all asserted
 claims (claims 1 and 9) of the ’083 patent. For the ’193 pa-
 tent, there were two accused products: Chromium Ge-
 nome/Exome and GemCode Long Read. The jury found
 both products infringe all asserted claims (claims 6 and 8)
 of the ’193 patent. For the ’407 patent, there were four ac-
 cused products: Chromium Genome/Exome; GemCode
 Long Read; Chromium Single Cell 3’; and Chromium Sin-
 gle Cell V(D)J. The jury found these products infringe the
 asserted claims (claims 1, 10, and 11).
                    C. Procedural History
     In February 2015, patent owner University of Chicago
 and its licensee RainDance Technologies, Inc.
 (“RainDance”) filed this patent infringement suit against
 10X. Bio-Rad subsequently purchased RainDance and, in
 May 2017, was substituted for RainDance in the litigation.
 After the litigation was filed, 10X modified its products to
 add 0.02% Kynar—a non-reactive amount of a fluorine-con-
 taining resin—to its microchannels. 10X concedes that the
 addition of this amount of Kynar is irrelevant to the func-
 tioning of its products. The district court held a jury trial
 in November 2018. The jury found the patents-in-suit not
 invalid and willfully infringed. The jury then awarded Bio-
 Rad damages in the amount of $23,930,716.
      10X moved for JMOL, asserting that the accused prod-
 ucts do not infringe, that its infringement was not willful,
 that the asserted claims are not valid, and that Bio-Rad
 failed to present a legally sufficient damages case. 10X also
 requested a remittitur and moved for a new trial. On July
 3, 2019, the district court denied 10X’s motion. Bio-Rad,
 for its part, moved for a permanent injunction, attorneys’
 fees, enhanced damages, supplemental damages, and pre-
 and post-judgment interest. On July 24, 2019, the district
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 8           BIO-RAD LABORATORIES, INC.   v. 10X GENOMICS INC.



 court granted the motion with respect to the permanent in-
 junction, supplemental damages, and pre- and post-judg-
 ment interest, but denied the motion with respect to
 attorneys’ fees and enhanced damages. This appeal fol-
 lowed. 2 We have jurisdiction pursuant to 28 U.S.C.
 §§ 1295(a)(1) and 1292(c)(2).
                       II. DISCUSSION
      We review a denial of JMOL or new trial under the law
 of the regional circuit. Energy Transp. Grp., Inc. v. William
 Demant Holding A/S, 697 F.3d 1342, 1350 (Fed. Cir. 2012).
 The Third Circuit “exercise[s] plenary review of an order
 granting or denying a motion for judgment as a matter of
 law and appl[ies] the same standard as the district court.”
 Lightning Lube, Inc. v. Witco Corp., 4 F.3d 1153, 1166 (3d
 Cir. 1993). JMOL is “granted only if, viewing the evidence
 in the light most favorable to the nonmovant and giving it
 the advantage of every fair and reasonable inference, there
 is insufficient evidence from which a jury reasonably could
 find” for the nonmovant. Id. The decision to grant or deny


     2    On August 19, 2019, 10X filed a Rule 8 Motion for
 a Stay Pending Appeal, seeking a stay of the district court’s
 injunction order. We initially ruled that 10X could con-
 tinue to sell its Linked-Reads and CNV products subject to
 the royalty and deposit requirements set forth in the dis-
 trict court’s injunction order. Order, Bio-Rad Labs., Inc. v.
 10X Genomics Inc., No. 2019-2255 (Fed. Cir. Aug. 19,
 2019). On September 24, 2019, we concluded that our prior
 stay order should remain in effect during the pendency of
 the appeal, noting that “10x Genomics indicates that, while
 it did not immediately have available to offer to new cus-
 tomers an instrument that would be capable of running
 only [the Linked-Reads and CNV] products, it could, in a
 matter of weeks, implement such a solution.” Order, Bio-
 Rad Labs., Inc. v. 10X Genomics Inc., No. 2019-2255, at *2
 (Fed. Cir. Sept. 24, 2019).
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 a new trial is committed to the discretion of the district
 court, which grants a new trial only where “a miscarriage
 of justice would result if the verdict were to stand” or where
 the verdict “shocks [the] conscience.” Williamson v. Consol.
 Rail Corp., 926 F.2d 1344, 1353 (3d Cir. 1991).
      On appeal, 10X argues that (1) it is entitled to JMOL
 of non-infringement of the three patents-in-suit; (2) the
 damages award should be vacated because it was based on
 both inadmissible and insufficient evidence; and (3) the dis-
 trict court abused its discretion in enjoining all five product
 lines. We address these arguments in turn.
             A. Infringement of the ’083 Patent
     The asserted claims of the ’083 patent recite “non-fluor-
 inated microchannels.” As of trial, 10X’s accused products
 contained microchannels with 0.02% Kynar—a fluorine-
 containing coating resin. The jury found that 10X’s ac-
 cused products, as modified, do not literally satisfy the
 “non-fluorinated microchannels” limitation but meet the
 limitation under the doctrine of equivalents. On appeal,
 10X argues that the district court erred in denying JMOL
 because “two independent legal principles” barred Bio-
 Rad’s theory of equivalence: prosecution history estoppel
 and claim vitiation. Appellant’s Br. 31–40. It also argues
 that its products cannot satisfy one of the claim limitations
 relating to the surface tension at the plug-fluid/carrier-
 fluid interface. As discussed below, we reject 10X’s argu-
 ments.
             1. Bio-Rad Was Not Estopped From
            Asserting the Doctrine of Equivalents
     “[E]quivalents remain a firmly entrenched part of the
 settled rights protected by the patent.” Festo Corp. v. Sho-
 ketsu Kinzoku Kogyo Kabushiki Co., 535 U.S. 722, 733
 (2002) (“Festo I”). There are certain limitations, however,
 on a patentee’s ability to obtain an infringement verdict
 under the doctrine of equivalents. One such limitation is
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 10             BIO-RAD LABORATORIES, INC.   v. 10X GENOMICS INC.



 prosecution history estoppel. See, e.g., id. at 737–40. An-
 other limitation—the doctrine of claim vitiation—ensures
 that “the application of the doctrine [of equivalents] . . . is
 not allowed such broad play as to effectively eliminate [a
 claim] element in its entirety.” Warner-Jenkinson Co. v.
 Hilton Davis Chem. Co., 520 U.S. 17, 29 (1997). Both are
 at issue in this appeal.
                a. Prosecution History Estoppel
     Prosecution history estoppel arises when a patent ap-
 plicant narrows the scope of his claims during prosecution
 for a reason “substantial[ly] . . . relating to patentability.”
 See generally Festo Corp. v. Shoketsu Kinzoku Kogyo Ka-
 bushiki Co., 344 F.3d 1359, 1366–67 (Fed. Cir. 2003) (en
 banc) (“Festo II”). A narrowing amendment is presumed to
 be a surrender of all equivalents within “the territory be-
 tween the original claim and the amended claim.” Festo I,
 535 U.S. at 740. This presumption can be overcome if the
 patentee can show that one of the following “exceptions” to
 prosecution history estoppel applies: (1) the rationale un-
 derlying the amendment bears no more than a tangential
 relation to the equivalent in question; (2) the equivalent
 was unforeseeable at the time of the application; or (3)
 there was some other reason suggesting that the patentee
 could not reasonably be expected to have described the
 equivalent. Id. at 740–41.
     “[W]hether prosecution history estoppel applies, and
 hence whether the doctrine of equivalents may be available
 for a particular claim limitation, presents a question of
 law.” Festo II, 344 F.3d at 1367–68. In making this deter-
 mination, we must “look to the specifics of the amendment
 and the rejection that provoked the amendment to deter-
 mine whether estoppel precludes the particular doctrine of
 equivalents argument being made.” Intervet Inc. v. Merial
 Ltd., 617 F.3d 1282, 1291 (Fed. Cir. 2010).
     The district court held that prosecution history estop-
 pel does not apply in this case because the amendment at
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 issue was only tangentially related to the accused equiva-
 lent. The court reasoned that, during prosecution, the in-
 ventors sought to distinguish the microchannels in their
 system from fluorinated prior art microchannels that
 would react with the carrier fluid. Bio-Rad, 396 F. Supp.
 3d at 377. In the court’s view, the objectively apparent rea-
 son for the amendment was to distinguish fluorinated mi-
 crochannels from microchannels that had no fluorinated
 properties. The district court concluded that an accused
 product like 10X’s—having minute or negligible quantities
 of fluorine that have no function in the product and do not
 react with the microchannels—could meet the “non-fluori-
 nated” limitation under the doctrine of equivalents. Id.
     On appeal, the parties do not dispute that the amend-
 ment at issue was narrowing, or that it was made to over-
 come prior art.      Instead, they dispute whether the
 tangentiality exception to prosecution history estoppel ap-
 plies. 10X argues, as it did before the district court, that
 prosecution history estoppel applies because the ’083 pa-
 tent inventors narrowed the claims to recite a “non-fluori-
 nated microchannel” to overcome Quake, which taught
 “fluorinated” microchannels. 10X argues that, with this
 amendment, the inventors surrendered all territory be-
 tween the original limitation—microchannels generally—
 and the amended limitation—non-fluorinated microchan-
 nels. Appellant’s Br. 33–34. Accordingly, 10X contends
 that the district court erred in applying the “tangentiality”
 exception for a narrowing amendment. 10X characterizes
 the district court’s analysis as “recasting” the inventors’
 disclaimer as covering only microchannels coated with flu-
 orine “for a purpose.” Id. at 35 (emphasis omitted). In
 10X’s view, by rewriting the claims during prosecution, the
 inventors surrendered the right to expand their monopoly
 to cover microchannels containing fluorine, “for whatever
 purpose.” Id. at 36.
     Bio-Rad argues that the tangentiality exception to
 prosecution history estoppel allows it to assert and prevail
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 12             BIO-RAD LABORATORIES, INC.   v. 10X GENOMICS INC.



 under the doctrine of equivalents. In Bio-Rad’s view, the
 reason for narrowing the claims was “peripheral, or not di-
 rectly relevant to the alleged equivalent.” Appellees’ Br. 20
 (quoting Festo II, 344 F.3d at 1369). Bio-Rad contends that
 the patentees amended the claims to make clear that the
 carrier fluid and the microchannel wall should be chemi-
 cally distinct, which bears no more than a tangential rela-
 tion to the alleged equivalent—microchannel walls
 containing a nominal amount of fluorine that is not chemi-
 cally distinct from the carrier fluid. We agree with Bio-
 Rad.
      The prosecution history of the ’083 patent establishes
 that the objectively apparent reason for adding the “non-
 fluorinated microchannels” limitation was no more than
 tangentially related to the equivalent at issue. In amend-
 ing the claims, the patentees sought to distinguish the
 claimed invention from Quake, which disclosed fluorinated
 microchannel wall coatings that would react with the car-
 rier fluid. The inventors argued that preventing droplets
 from sticking to the walls of the microchannels requires the
 surfactant to be chemically similar to the carrier fluid and
 chemically different from the channel walls. The inventors
 therefore amended the claims to make clear that the car-
 rier fluid and the microchannel wall should be chemically
 distinct. By claiming non-fluorinated microchannels and a
 fluorinated surfactant, the inventors made sure that, in
 contrast to the Quake disclosure, the carrier fluid and mi-
 crochannels in the claimed invention would not react with
 each other, thereby preventing droplets from sticking to
 the walls of the microchannels. As such, the inventors’ de-
 cision to add the “non-fluorinated microchannels” limita-
 tion must be considered in the context of adding, at the
 same time, the limitation of a “fluorinated surfactant” to
 the carrier fluid.
     The inventors’ statements during prosecution confirm
 that the “rationale underlying the [narrowing] amendment
 [bore] no more than a tangential relation to the equivalent
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 in question”—here, microchannels containing negligible
 amounts of fluorine, which cannot react with the carrier
 fluid. See Festo I, 535 U.S. at 740–41. As the district court
 explained, the inventors surrendered microchannels
 coated with fluorine “for a purpose—not those containing
 de minimis amounts of fluorine that have no effect on how
 the microchannel functions in the system.” Bio-Rad, 396
 F. Supp. 3d at 377 (emphasis added). As such, Bio-Rad was
 not barred from asserting that microchannels containing
 negligible amounts of fluorine are equivalent to “non-fluor-
 inated microchannels.”
      We reject 10X’s argument that the “Quake patent un-
 ambiguously contained the accused equivalent,” and there-
 fore, this amendment cannot be tangential. Appellant’s Br.
 34–35. The crux of the tangentiality inquiry remains “the
 patentee’s objectively apparent reason for the narrowing
 amendment . . . [as] discernible from the prosecution his-
 tory record.” Festo II, 344 F.3d at 1369. As explained
 above, the prosecution history record reveals that the rea-
 son for the amendment was to distinguish microchannels
 that reacted with carrier fluids. Quake disclosed fluori-
 nated microchannels generally. It did not expressly dis-
 close microchannels with non-reacting, negligible levels of
 fluorine, like in the accused equivalent. The question here
 is not whether Quake disclosed fluorinated microchannels,
 but rather, whether Quake taught the use of non-reactive
 amounts of fluorination in the microchannels. It did not.
 Accordingly, the narrowing amendment can only be said to
 have a tangential relation to the equivalent at issue—neg-
 ligibly fluorinated microchannels, or, put differently, mi-
 crochannels with non-fluorinated properties.
      The parties each cite several of our cases in support of
 or against the application of the tangentiality exception to
 prosecution history estoppel. Prosecution history estoppel,
 including the tangentiality inquiry, is always a case-spe-
 cific analysis. The objectively apparent reason discernable
 from the prosecution history record will, accordingly, differ
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 14             BIO-RAD LABORATORIES, INC.   v. 10X GENOMICS INC.



 in each case. For example, we recently considered these
 issues in Amgen Inc. v. Amneal Pharmaceuticals LLC, 945
 F.3d 1368 (Fed. Cir. 2020). There, the accused product
 used “pregelatinized starch” as a binder, and the asserted
 claim did not list pregelatinized starch in its Markush
 group reciting binders. Id. at 1380. The patent owner as-
 serted infringement under the doctrine of equivalents, ar-
 guing that pregelatinized starch functioned as a binder in
 the accused product. We noted that the patent owner re-
 vised the claim’s binder limitations to be in Markush group
 format to overcome prior art references that taught the use
 of pregelatinized starch as a binder. Id. at 1382. Accord-
 ingly, we concluded that the amendment—made to avoid
 prior art that contains the equivalent in question—was not
 tangential. Id. Because the prior art references at issue
 taught the use of the alleged equivalent for the claimed
 function, the tangentiality exception to prosecution history
 estoppel could not apply. Here, by contrast, Quake did not
 teach the use of the alleged equivalent—negligibly fluori-
 nated microchannels or those with no fluorinated proper-
 ties.
     We also recently addressed the tangentiality exception
 in Eli Lilly v. Hospira, Inc., where the patent owner nar-
 rowed the claims during prosecution to recite “pemetrexed
 disodium” instead of “an antifolate.” 933 F.3d 1320,
 1325–26 (Fed. Cir. 2019). The accused equivalent at issue
 was pemetrexed ditromethamine, which is functionally
 identical to pemetrexed disodium. Id. at 1327. We con-
 cluded that “[t]he reason for Lilly’s amendment . . . was to
 narrow original claim 2 to avoid Arsenyan, which only dis-
 closes treatments using methotrexate, a different antifo-
 late.” Id. at 1331. Thus, claiming the functionally
 equivalent pemetrexed salts was tangential to overcoming
 prior art disclosing an antifolate other than pemetrexed.
 Here too, functionally equivalent microchannels (i.e., mi-
 crochannels with no fluorinated properties) are tangential
 to the patentees’ reason for distinguishing Quake, which
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 BIO-RAD LABORATORIES, INC.   v. 10X GENOMICS INC.          15



 disclosed fluorinated microchannels. We find this case
 more analogous to Eli Lilly than to Amgen.
     Accordingly, we conclude that the district court cor-
 rectly held that prosecution history estoppel does not apply
 in this case.
                      b. Claim Vitiation
     Claim vitiation presents another bar to a finding of in-
 fringement under the doctrine of equivalents. “[S]aying
 that a claim element would be vitiated is akin to saying
 that there is no equivalent to the claim element in the ac-
 cused device based on the well-established ‘function-way-
 result’ or ‘insubstantial differences’ tests.” Brilliant In-
 struments, Inc. v. GuideTech, LLC, 707 F.3d 1342, 1347
 (Fed. Cir. 2013). More recently, we have explained that vi-
 tiation “is not an exception or threshold determination that
 forecloses resort to the doctrine of equivalents, but is in-
 stead a legal conclusion of a lack of equivalence based on
 the evidence presented and the theory of equivalence as-
 serted.” UCB, Inc. v. Watson Labs., Inc., 927 F.3d 1272,
 1283 (Fed. Cir. 2019) (quoting Cadence Pharm. Inc. v. Ex-
 ela PharmSci Inc., 780 F.3d 1364, 1371 (Fed. Cir. 2015));
 see also DePuy Spine, Inc. v. Medtronic Sofamor Danek,
 Inc., 469 F.3d 1005, 1017 (Fed. Cir. 2006) (“[T]he ‘all ele-
 ments’ rule generally is not met—and therefore a claim
 limitation can be said to be vitiated—if the theory or evi-
 dence of equivalence is legally incapable of establishing
 that the differences between the limitation in the claim and
 the accused device are insubstantial; i.e., if the theory or
 evidence is so legally insufficient as to warrant a holding of
 non-infringement as a matter of law.”).
     On appeal, 10X argues that the doctrine of equivalents
 is unavailable to Bio-Rad because “fluorinated” and “non-
 fluorinated” are “diametric opposites” and because a fluor-
 inated microchannel is the “antitheses” of a non-fluori-
 nated microchannel. Appellant’s Br. 37–40 (citing, e.g.,
 Moore U.S.A., Inc. v. Standard Register Co., 229 F.3d 1091,
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 16             BIO-RAD LABORATORIES, INC.   v. 10X GENOMICS INC.



 1115 n.5 (Fed. Cir. 2000)). In 10X’s view, allowing Bio-Rad
 to argue that fluorinated microchannels are equivalent to
 non-fluorinated microchannels entirely vitiates the “non-
 fluorinated microchannel” limitation. 10X also cites sev-
 eral of our prior cases where we found that a claim element
 cannot be supplied by an alleged equivalent that was the
 opposite of the missing element. Despite some surface ap-
 peal, these arguments do not hold up under even minimal
 scrutiny.
     10X attempts to extend our cases regarding claim viti-
 ation beyond their facts. Relying on Deere & Co. v. Bush
 Hog, LLC, 703 F.3d 1349, 1356 (Fed. Cir. 2012), the district
 court concluded that a reasonable jury could find on the
 facts presented here that non-fluorinated microchannels
 and minimally-fluorinated microchannels with no reactive
 properties are equivalent. 10X criticizes the district court’s
 reliance on Deere as “invok[ing] only dicta.” Appellant’s
 Br. 39. It also argues that Deere did not override our prior
 cases discussing “opposites” but instead dealt with a sce-
 nario where the claim element did not present a binary
 choice. Appellant’s Reply Br. 7. 10X, however, ignores the
 fact that we have repeatedly emphasized the principle out-
 lined in Deere in subsequent cases. See, e.g., Cadence
 Pharm. Inc., 780 F.3d at 1371 (collecting cases).
     In Brilliant Instruments, for example, we explained
 that vitiation comes into play when the alleged equivalent
 is “diametrically opposed” to the missing claim element.
 “[W]hen the accused structure has an element that is the
 opposite of the claimed element,” it is “more difficult” for a
 patentee to succeed on a theory of equivalents. 707 F.3d
 at 1347. We also explained that “[i]f the claimed and ac-
 cused elements are recognized by those of skill in the art to
 be opposing ways of doing something, they are likely not
 insubstantially different.” Id. at 1347–48. And, we noted,
 “this concept [applies] to cases where we have recognized
 that two alternatives exist that are very different from
 each other and therefore cannot be equivalents for
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 BIO-RAD LABORATORIES, INC.   v. 10X GENOMICS INC.          17



 infringement purposes.” Id. at 1348 (citing, e.g., Moore
 U.S.A., Inc., 229 F.3d at 1106).
     Notably, in Cadence Pharmaceuticals, we cautioned
 against using labels like “antithesis” in lieu of conducting
 the proper inquiry of infringement under the doctrine of
 equivalents:
    Characterizing an element of an accused product as
    the “antithesis” of a claimed element is also a con-
    clusion that should not be used to overlook the fac-
    tual analysis required to establish whether the
    differences between a claimed limitation and an ac-
    cused structure or step are substantial vel non. The
    determination of equivalence depends not on labels
    like “vitiation” and “antithesis” but on the proper
    assessment of the language of the claimed limita-
    tion and the substantiality of whatever relevant dif-
    ferences may exist in the accused structure.
 Cadence Pharm., 780 F.3d at 1372.
      Accordingly, we reject 10X’s attempt to limit the in-
 quiry to a binary choice between “fluorinated” and “non-
 fluorinated” microchannels, and its conclusion that in-
 fringement under the doctrine of equivalents is unavaila-
 ble to Bio-Rad as a matter of law. The appropriate inquiry
 is whether a reasonable juror could have found that a neg-
 ligibly-fluorinated microchannel performs the same func-
 tion, in the same way, and achieves the same result, as a
 non-fluorinated microchannel. Here, based on the evidence
 presented at trial—including the testimony of Bio-Rad’s ex-
 pert, Dr. Sia—the district court concluded that a reasona-
 ble juror could find that a 0.02% Kynar-containing
 microchannel is insubstantially different from a non-fluor-
 inated microchannel. Bio-Rad, 396 F. Supp. 3d at 378. The
 non-fluorinated microchannel claim limitation is not
 stripped of meaning, or “effectively eliminate[d],” Warner-
 Jenkinson, 520 U.S. at 29, by Bio-Rad’s theory and the
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 18             BIO-RAD LABORATORIES, INC.   v. 10X GENOMICS INC.



 jury’s finding that the accused microchannel, having too lit-
 tle fluorine to alter its reactive properties, is an equivalent.
     10X does not challenge that substantial evidence sup-
 ports the jury’s finding that a 0.02% Kynar-containing mi-
 crochannel is insubstantially different from a non-
 fluorinated microchannel. Accordingly, 10X’s challenges
 concerning the infringement under the doctrine of equiva-
 lents fail.
      2. Substantial Evidence Supports the Jury’s Verdict
        that 10X’s Accused Products Meet the Claimed
                Surface Tension Relationship
     10X next argues that it is entitled to JMOL of non-in-
 fringement of the ’083 patent because none of its products
 have a plug-fluid/microchannel wall interface. Appellant’s
 Br. 40–42. According to 10X, without such an interface, its
 products cannot satisfy the claim limitation “wherein the
 fluorinated surfactant is present at a concentration such
 that the surface tension at the plug-fluid/microchannel
 wall interface is higher than the surface tension at the
 plug-fluid/carrier-fluid interface.” Id. at 40 (quoting ’083
 patent, col. 73 ll. 16–21) (emphasis omitted). 10X contends
 that the droplets in its chips are fully encased by the car-
 rier fluid and do not touch the channel wall, meaning they
 do not have the plug-fluid/microchannel wall interface, and
 thus cannot infringe.
     Bio-Rad responds that the claims do not require the
 plug-fluid to be in actual physical contact with the micro-
 channel walls. Appellees’ Br. 24. In Bio-Rad’s view, the
 asserted claims merely require that the droplet/wall sur-
 face tension is higher than the droplet/carrier-fluid surface
 tension. According to Bio-Rad, the purpose of the claimed
 surface tension relationship is to allow the formation of
 plugs/droplets that do not stick to the channel walls, just
 like those in 10X’s products. Id. at 25. We again agree with
 Bio-Rad.
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 BIO-RAD LABORATORIES, INC.   v. 10X GENOMICS INC.             19



      The district court correctly noted that the claims do not
 require direct contact between the plug-fluid and the mi-
 crochannel wall. They only require that the surface tension
 at the plug-fluid/microchannel wall interface is higher than
 that between the carrier fluid and the plug fluid. The spec-
 ification explains the purpose of this claimed surface ten-
 sion relationship: “If this condition is not satisfied, plugs
 tend to adhere to the channel walls and do not undergo
 smooth transport.” ’083 patent, col. 20 ll. 56–58. And, the
 specification clearly sets forth the purpose of introducing
 the surfactant:
    Because the walls of the channels (PDMS, not fluor-
    inated) and the carrier-fluid (fluorinated oil) are
    substantially different chemically, when a fluori-
    nated surfactant is introduced, the surfactant re-
    duces the surface tension at the oil-water interface
    preferentially over the wall-water interface. This
    allows the formation of plugs that do not stick to the
    channel walls.
 Id. at col. 20 l. 63–col. 21 l. 2. Given this, we find that 10X’s
 argument that its accused products cannot meet this limi-
 tation is without merit.
     We also agree with the district court that sufficient ev-
 idence supports the jury’s finding that 10X’s products meet
 the claimed surface tension limitation. 10X’s own expert,
 Dr. Huck, admitted that the surface tension relationship is
 met if the droplets do not contact channel walls. The par-
 ties do not dispute that this is the case for 10X’s accused
 products. The jury also considered the testimony of Bio-
 Rad’s expert, Dr. Sia, who presented testing evidence
 demonstrating that the claim limitation was met. Accord-
 ingly, we conclude that the district court properly denied
 10X’s motion for JMOL of non-infringement of the ’083 pa-
 tent and affirm the judgment of infringement of this pa-
 tent.
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 20             BIO-RAD LABORATORIES, INC.   v. 10X GENOMICS INC.



        B. Infringement of the ’407 and ’193 Patents
      10X’s non-infringement arguments regarding the ’407
 and ’193 patents are based on its contention that the dis-
 trict court misconstrued the asserted claims of these pa-
 tents. Specifically, 10X argues that, contrary to the district
 court’s construction, the preambles of these patents’ inde-
 pendent claims are limiting. We review claim construction
 de novo, reviewing subsidiary factual findings based on ex-
 trinsic evidence for clear error. Teva Pharm. USA, Inc. v.
 Sandoz, Inc., 574 U.S. 318 (2015).
      Whether a preamble is limiting is “determined on the
 facts of each case in light of the overall form of the claim,
 and the invention as described in the specification and il-
 luminated in the prosecution history.” Applied Materials,
 Inc. v. Advanced Semiconductor Materials Am., Inc., 98
 F.3d 1563, 1572–73 (Fed. Cir. 1996). A preamble limits the
 claimed invention if it recites essential structure or steps,
 or if it is “necessary to give life, meaning, and vitality” to
 the claim. Pitney Bowes, Inc. v. Hewlett-Packard Co., 182
 F.3d 1298, 1305 (Fed. Cir. 1999). If the claim uses the pre-
 amble only to state a purpose or intended use for the inven-
 tion, then the preamble is not limiting. Catalina Mktg.
 Int’l, Inc. v. Coolsavings.com, Inc., 289 F.3d 801, 808 (Fed.
 Cir. 2002). And, a preamble is generally not limiting un-
 less there is “clear reliance on the preamble during prose-
 cution to distinguish the claimed invention from the prior
 art.” Id. Reliance on a preamble phrase for antecedent ba-
 sis, however, may limit claim scope. Bell Commc’ns Re-
 search, Inc. v. Vitalink Commc’ns Corp., 55 F.3d 615, 620
 (Fed. Cir. 1995).
     The preamble at issue recites “[a] method for conduct-
 ing a reaction in plugs in a microfluidic system, comprising
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 BIO-RAD LABORATORIES, INC.   v. 10X GENOMICS INC.          21



 the steps of . . . .” ’407 patent, col. 78 ll. 54–55. 3 At the
 Markman stage, the district court found that the preamble
 was limiting “only to the extent that it provides an ante-
 cedent basis for the terms ‘microfluidic system’ and ‘reac-
 tion.’” Raindance Techs., Inc. v. 10X Genomics, Inc., No.
 1:15-CV-00152-RGA, 2017 WL 382235, at *7 (D. Del. Jan.
 26, 2017)). It found that the preamble at issue “states an
 intended use for the invention, ‘followed by the body of the
 claim, in which the claim limitations describing the inven-
 tion are recited.’” Id. (quoting TomTom, Inc. v. Adolph,
 790 F.3d 1315, 1324 (Fed. Cir. 2015)). It also found that
 the invention, as claimed, was “‘structurally complete’
 without the remaining preamble language.” Id.
      In its motion for JMOL, 10X argued that, under the
 correct claim construction, the preambles limit the claims
 to methods of conducting reactions inside a microfluidic
 system, i.e., to “on-chip” reactions only. The district court
 rejected 10X’s argument. It found that the preamble terms
 “reaction” and “microfluidic systems” provide antecedent
 basis for the use of those terms in the body of the claim, but
 that this does not necessarily convert the entire preamble
 into a limitation. Bio-Rad, 396 F. Supp. 3d at 380 (“While
 portions of a preamble may be limiting where those por-
 tions provide an antecedent basis for terms appearing in
 the body of the claim, it is inappropriate to construe an en-
 tire preamble as limiting if the rest of the preamble lan-
 guage is not limiting.”). According to the district court, the
 italicized portion of the preamble “conducting a reaction in
 plugs in a microfluidic system” was not limiting because



     3   The preamble of the claims of the ’193 patent recite
 an “autocatalytic reaction” instead of a “reaction.” ’193 pa-
 tent, col. 78 ll. 8–9. 10X states that the same arguments
 apply to the preambles of both the ’407 and ’193 patent
 claims and Bio-Rad offers no dispute on that point. Accord-
 ingly, our analysis applies to both patents.
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 22             BIO-RAD LABORATORIES, INC.   v. 10X GENOMICS INC.



 this language does not “provide an antecedent basis for the
 rest of the claim and follows the standard pattern of a
 ‘method for a purpose or intended use comprising,’ followed
 by the body of the claim.” Id. at 381 (citing TomTom, 790
 F.3d at 1324).
     On appeal, 10X again argues that the preamble term
 “reaction in plugs in a microfluidic system” is limiting. In
 10X’s view, the preamble requires the chemical reaction to
 take place in the droplets while the droplets are inside the
 microfluidic system, i.e., the claimed methods are limited
 to “on-chip” reactions. By contrast, 10X argues, the reac-
 tions in its accused products are “off-chip” reactions—they
 are not completed until after a researcher removes the
 droplets from the microchannel and places them in a ther-
 mal cycler. Appellant’s Br. 51–53.
      10X argues that, as the district court found, the pream-
 ble clearly provides antecedent basis for the claim limita-
 tions “the microfluidic system” and “the reaction.” It
 contends that, taken in conjunction with the specification
 and the prosecution history of the ’407 patent, this estab-
 lishes the drafter’s intent to treat the entire preamble as
 limiting. Although we held in TomTom that it is possible
 for one part of the preamble to be limiting even though an-
 other portion is not, 10X asserts that TomTom does not ap-
 ply where, as here, the preamble terms at issue were not
 distinct phrases. According to 10X, it was error to give lim-
 iting effect to certain terms in the preamble (“reaction” and
 “microfluidic system”) but not to other terms surrounding
 those limiting parts (“conducting” and “in”).
     In Bio-Rad’s view, the district court correctly found
 that the disputed language is a non-limiting statement of
 intended use or purpose. Bio-Rad also argues that, under
 TomTom, an entire preamble need not be limiting simply
 because it provides an antecedent basis in part. Bio-Rad
 further argues that 10X’s non-enablement theory at trial
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 BIO-RAD LABORATORIES, INC.   v. 10X GENOMICS INC.         23



 contradicts its arguments here. This time, we agree with
 10X.
      The district court’s application of TomTom to the facts
 before it is erroneous. In TomTom, we held that the district
 court erred in determining that it had to construe the en-
 tire preamble if it construed a portion of it. TomTom, 790
 F.3d at 1322–24. The two-part preamble of the asserted
 claim recited: “[1] [a] method for generating and updating
 data [2] for use in a destination tracking system of at least
 one mobile unit comprising . . . .” Id. at 1322 (alteration
 omitted). We held that the first part of the preamble,
 “method for generating and updating data,” was not limit-
 ing and did not provide an antecedent basis for any claim
 terms. Id. at 1323–24. We also found that the term did not
 recite essential structure or steps, or give necessary life,
 meaning, and vitality to the claim; rather, it stated “a pur-
 pose or intended use.” Id. At the same time, we held that
 the second part of the preamble, “destination tracking sys-
 tem of at least one mobile unit,” was limiting because it
 provided antecedent basis for “the mobile unit” recited in
 the body of the claims. Id. Thus, we found that TomTom
 involved a partially-limiting preamble.
     Crucially, unlike TomTom, the preamble in this case
 cannot be neatly packaged into two separate portions. Nor
 does it simply recite a method for an intended use or pur-
 pose. The district court held that the preamble terms “re-
 action” and “microfluidic systems” provide antecedent
 basis for the use of those terms in the body of the claim.
 We agree with the court on this point. But we disagree that
 these limiting terms can be read separately from the re-
 mainder of the preamble. The language relied upon for an-
 tecedent basis in the preamble at issue is intertwined with
 the rest of the preamble. The term “conducting” in the pre-
 amble is not analogous to the non-limiting language at is-
 sue in TomTom.
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 24             BIO-RAD LABORATORIES, INC.   v. 10X GENOMICS INC.



     We also are disinclined to sanction finding a preamble
 “partially” limiting by splicing it as the district court did
 here. The fact that the terms “reaction” and “microfluidic
 systems” provide antecedent basis for these terms in the
 body of the claim is a strong indication that the preamble
 acts “as a necessary component of the claimed invention.”
 Eaton Corp. v. Rockwell Int’l Corp., 323 F.3d 1332, 1339
 (Fed. Cir. 2003). Based on the antecedent relationship, it
 is clear the claim drafters intended to limit the claimed
 methods to on-chip reactions, using both the preamble and
 the body of the claim to define the claimed invention. 4 Ac-
 cordingly, we conclude that the district court’s claim con-
 struction is erroneous. Under the correct construction, the
 claimed methods are limited to on-chip reactions.
     Bio-Rad argues that, even if the preamble is limiting,
 the jury’s infringement verdict should be upheld. As a
 court of review, we refuse to decide, in the first instance,
 whether 10X’s systems would infringe under the correct
 construction. Even Bio-Rad concedes that the district court
 precluded 10X from arguing that the entire preamble is
 limiting. Accordingly, we vacate the district court’s judg-
 ment of infringement of the claims of the ’407 and ’193 pa-
 tents and remand for a new trial on those issues.




      4  The prosecution history of the patent, although not
 dispositive in this case, provides additional support for con-
 struing the claims as limited to reactions in a microfluidic
 system. During prosecution, the examiner amended both
 the preambles and the titles of the patents to specify that
 the reactions are conducted “in plugs in the microfluidic
 system.” J.A. 8625–29; J.A. 8632.
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 BIO-RAD LABORATORIES, INC.   v. 10X GENOMICS INC.          25



                         C. DAMAGES

      Despite vacating the district court’s judgment of in-
 fringement of two of the patents-in-suit, we proceed with
 considering the parties’ arguments concerning damages be-
 cause we affirm the judgment of infringement of the ’083
 patent—which covers all six accused product lines. The
 jury verdict and jury instructions show that the damages
 award is not predicated on infringement of any one patent.
 J.A. 378 (“If you found that 10X Genomics infringed any of
 the asserted claims of the ’083, ’193, or ’407 [p]atents . . .
 then with respect to that claim or those claims, please an-
 swer [the question on damages].”); J.A. 410 (“If you find
 that Plaintiffs have established infringement of a valid pa-
 tent claim of the patents-in-suit, Plaintiffs will be entitled
 to a reasonable royalty to compensate them for that in-
 fringement.”). As Bio-Rad explained during oral argument,
 affirming the judgment of infringement on the ’083 pa-
 tent—which includes the only asserted apparatus claims—
 would leave the damages award undisturbed. Oral Arg. at
 21:18–44, available at http://oralarguments.cafc.uscourts.g
 ov/default.aspx?fl=19-2255.mp3. 10X did not dispute this
 point either at oral argument or in its briefing to us. In
 fact, in its opening brief, 10X argued that reversal as to the
 ’083 patent would affect the damages award, see Appel-
 lant’s Br. 42, but did not make such an argument for the
 ’407 and ’193 patents.
      The jury awarded almost $24 million in damages—the
 full requested amount based on a reasonable royalty rate
 of 15%. On appeal, 10X argues that the damages award
 should be vacated because Bio-Rad’s expert relied on li-
 censes that were not comparable to the hypothetical nego-
 tiation. 10X further argues that Bio-Rad’s expert did not
 apportion damages to the value of the patented technology.
 As discussed below, we reject these arguments.
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 26             BIO-RAD LABORATORIES, INC.   v. 10X GENOMICS INC.



      1. The District Court Did Not Abuse Its Discretion
         in Allowing the Jury to Consider Testimony
            Regarding the Three Licenses at Issue
     At trial, the parties used the hypothetical negotiation
 or the “willing licensor-willing licensee” approach for calcu-
 lating reasonable royalty damages. This approach at-
 tempts to calculate the royalty rate the parties would have
 agreed upon had they negotiated an agreement prior to the
 start of the infringement. In determining a reasonable roy-
 alty, parties frequently rely on comparable license agree-
 ments. See Georgia–Pacific Corp. v. U.S. Plywood Corp.,
 318 F. Supp. 1116, 1120 (S.D.N.Y. 1970); see also Radio
 Steel & Mfg. Co. v. MTD Prods., Inc., 788 F.2d 1554, 1557
 (Fed. Cir. 1986) (“The determination of a reasonabl[e] roy-
 alty . . . is based . . . on the royalty to which a willing licen-
 sor and a willing licensee would have agreed at the time
 the infringement began.”). Assessing the comparability of
 licenses requires a consideration of whether the license at
 issue involves comparable technology, is economically com-
 parable, and arises under comparable circumstances as the
 hypothetical negotiation. See generally LaserDynamics,
 Inc. v. Quanta Computer, Inc., 694 F.3d 51 (Fed. Cir. 2012).
     We review the district court’s decision to admit expert
 testimony for abuse of discretion. Gen. Elec. Co. v. Joiner,
 522 U.S. 136, 146 (1997). The jury’s determination of the
 amount of damages is an issue of fact, which we review for
 substantial evidence. Lucent Techs., Inc. v. Gateway, Inc.,
 580 F.3d 1301, 1310 (Fed. Cir. 2009). A jury’s damages
 award “must be upheld unless the amount is grossly exces-
 sive or monstrous, clearly not supported by the evidence,
 or based only on speculation or guesswork.” Id. (internal
 quotation marks omitted).
     At trial, Bio-Rad’s damages expert, Mr. Malackowski,
 based his reasonable royalty calculation on three licenses
 that he deemed comparable: (1) the Caliper/RainDance li-
 cense (2) the Applera/Bio-Rad license, and (3) the Applied
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 BIO-RAD LABORATORIES, INC.   v. 10X GENOMICS INC.         27



 Bio/QuantaLife license. In its motion for JMOL, 10X ar-
 gued that Mr. Malackowski’s testimony was not suffi-
 ciently tied to the facts of the case because it was based on
 technologically noncomparable licenses. The district court
 agreed with 10X as to the Applera/Bio-Rad license, finding
 that Bio-Rad had failed to present sufficient evidence of the
 technological comparability of this license. But it nonethe-
 less denied 10X’s motion for JMOL because the other two
 licenses provided sufficient support for Mr. Malackowski’s
 reasonable royalty opinions. Bio-Rad, 396 F. Supp. 3d at
 386.
     On appeal, 10X argues that Mr. Malackowski’s expert
 opinion should be vacated because it was based on evidence
 that was “both inadmissible and insufficient.” Appellant’s
 Br. 54. 10X also provides extensive argument as to why
 each of the three licenses is not technologically comparable
 to the technology at issue in the hypothetical negotiation.
 Id. at 59–64.
     This court has often excluded licenses that are techno-
 logically or economically non-comparable. See, e.g., La-
 serDynamics, 694 F.3d at 77–78 (“The propriety of using
 prior settlement agreements to prove the amount of a rea-
 sonable royalty is questionable.”); ResQNet.com, Inc. v.
 Lansa, Inc., 594 F.3d 860, 873 (Fed. Cir. 2010) (district
 court erred by considering certain licenses and adjusting
 “upward” the reasonable royalty rate “without any factual
 findings that accounted for the technological and economic
 differences between those licenses”). The court has also
 held, however, that the issue of comparability is often one
 of sufficiency of the evidence, not admissibility. Ericsson,
 Inc. v. D-Link Sys., Inc., 773 F.3d 1201, 1227 (Fed. Cir.
 2014) (“[T]he fact that a license is not perfectly analogous
 generally goes to the weight of the evidence, not its admis-
 sibility.”); see also Finjan, Inc. v. Secure Computing Corp.,
 626 F.3d 1197, 1211 (Fed. Cir. 2010); ActiveVideo Net-
 works, Inc. v. Verizon Commc’ns, Inc., 694 F.3d 1312, 1333
 (Fed. Cir. 2012). Finjan, for example, involved a license
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 28             BIO-RAD LABORATORIES, INC.   v. 10X GENOMICS INC.



 which related to a lump sum payment rather than a run-
 ning royalty. 626 F.3d at 1211–12. We affirmed a damages
 award because the “differences permitted the jury to
 properly discount [that license].” Id. at 1212. Likewise, in
 ActiveVideo, the damages expert relied on two agreements,
 one of which did not involve the patents or technologies in
 the case. 694 F.3d at 1333. We concluded that the district
 court did not abuse its discretion by failing to exclude the
 testimony of the damages expert because the “degree of
 comparability” of the license agreements is a “factual is-
 sue[] best addressed by cross examination and not by ex-
 clusion.” Id.
     Here, the district court concluded that Mr. Malackow-
 ski had met a showing of “baseline comparability” and that
 the “degree of comparability is a factual issue best ad-
 dressed through cross examination.” Bio-Rad, 396 F.
 Supp. 3d at 388. The Caliper/RainDance and Applied-
 Bio/QuantaLife licenses covered patents related to micro-
 fluids. The third license, Applera/Bio-Rad, dealt with
 thermal PCR cyclers—instruments that are used in bio-
 chemical reactions. The “degree of comparability” was ap-
 propriately left for the jury to decide. See Active Video, 694
 F.3d at 1333. Accordingly, we see no abuse of discretion in
 allowing Mr. Malackowski to testify about these licenses.
     We are also not persuaded by 10X’s argument that, be-
 cause the district court ultimately concluded that Bio-Rad
 had not presented sufficient evidence of comparability of
 the Applera/Bio-Rad agreement, the jury should never
 have heard testimony regarding this agreement. 10X con-
 tends that it is entitled to a new trial on this ground alone,
 unless it is “highly probable” that the error of admitting
 testimony about this license did not affect the jury’s ver-
 dict. Appellant’s Br. 58 (quoting Hirst v. Inverness Hotel
 Corp., 544 F.3d 221, 228 (3d Cir. 2008)). We see several
 problems with 10X’s argument.
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 BIO-RAD LABORATORIES, INC.   v. 10X GENOMICS INC.          29



      Even assuming 10X sufficiently developed this argu-
 ment in its opening appellate brief (which it did not), 10X
 does not challenge the jury instructions regarding the cal-
 culation of a reasonable royalty, which properly instructed
 the jury to account for any differences between the licenses.
 J.A. 415 (“[I]f you choose to rely upon evidence from any
 other license agreements, you must account for any differ-
 ences between those licenses and the hypothetically nego-
 tiated license . . . in terms of the technologies and economic
 circumstances of the contracting parties.”). The record also
 shows that 10X’s Daubert motion regarding Mr. Malackow-
 ski’s opinions was directed to his overall testimony, not to
 the exclusion of any one agreement. 10X also did not move
 the court during trial to exclude the specific agreement.
 10X’s arguments on this issue again conflate the question
 of admissibility with the question of degree of comparabil-
 ity of the licenses. On this record, we see no abuse of dis-
 cretion in admitting Mr. Malackowski’s testimony, even if
 the district court ultimately determined that Bio-Rad did
 not provide substantial evidence of technological compara-
 bility of one of the three licenses. Accordingly, we disagree
 with 10X that the jury should have never heard testimony
 regarding the Applera/Bio-Rad license, and reject its re-
 quest for a new trial.
        2. The Jury’s Damages Award Is Supported
                 by Substantial Evidence
     10X also argues that no reasonable juror could find the
 three licenses comparable to the claimed invention. In-
 stead of the three licenses relied upon by Mr. Malackowski,
 10X contends that the University of Chicago/RainDance li-
 cense is the most comparable license because it concerns
 the patents-in-suit. We are not persuaded.
     The evidence at trial addressed each of the non-compa-
 rability arguments 10X is now raising on appeal. As to the
 University of Chicago/RainDance license, Mr. Malackow-
 ski opined that this license, from a university to a licensor
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 30             BIO-RAD LABORATORIES, INC.   v. 10X GENOMICS INC.



 in the nascent period of the droplet technology, is not com-
 parable to the hypothetical negotiation between two com-
 petitors in the context of a more developed field. He also
 explained that it was a non-competitive and exclusive li-
 cense, further distinguishing it from the hypothetical nego-
 tiation. 10X then cross-examined Mr. Malackowski on this
 issue. The jury was free to accept this testimony and to
 reject the 1–3% royalty rate proposed by 10X based on the
 University of Chicago/RainDance agreement.
     As to the comparability of the Caliper/RainDance li-
 cense, 10X argues that the license is not comparable for
 several reasons: (1) it deals with consumables (reagents
 and chips) and not with expensive instruments; (2) the li-
 cense involved 500+ patents relating to microfluids, not
 three patents dealing with specific droplet generation and
 manipulation; and (3) the 15% rate was “pure fiction” and
 never really actualized because RainDance and Caliper
 never competed in the licensed space. Appellant’s Br.
 62–63.
      The evidence at trial was sufficient to show compara-
 bility of the Caliper/RainDance agreement. The jury heard
 the testimony of Bio-Rad’s technical expert, Dr. Sia, who
 testified that the Caliper patents were comparable because
 they dealt with microfluids and the asserted patents also
 deal with the same subject matter, but specifically with
 droplets. Bio-Rad’s corporate witness, Ms. Tumolo, also ac-
 counted for the difference in the number of patents in the
 Caliper/RainDance agreement and the hypothetical nego-
 tiation. According to Ms. Tumolo, the large number of Cal-
 iper patents in the Caliper/RainDance license were a
 necessary but very small part of the licensed RainDance
 portfolio; the main technology was the droplet technology,
 covered by the three patents-in-suit. Ms. Tumolo also ad-
 dressed the 15% royalty rate, confirming that once
 RainDance directly competed with Caliper, the 15% royalty
 rate would apply. As to differentiating between licenses
 relating to consumables versus instruments, most of the
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 reasonable royalty damages at issue come from sales of
 consumables, not instruments. Accordingly, a reasonable
 juror could have concluded that the Caliper/RainDance li-
 cense was comparable to the hypothetical negotiation, and
 substantial evidence supports the jury’s verdict that a 15%
 reasonable royalty applies.
      10X makes similar arguments regarding the Applied-
 Bio/QuantaLife license. It argues that (1) the license does
 not require payments for instruments or even chips, and
 the 10–15% royalty rate for reagents cannot be directly
 translated to a royalty rate for expensive instruments; (2)
 the license focuses on a PCR enzyme that transformed the
 field; and (3) Mr. Malackowski did not address these differ-
 ences. Appellant’s Br. 60–61. We disagree.
      The evidence at trial supports a finding of comparabil-
 ity for this agreement as well. For example, the jury con-
 sidered the testimony of Dr. Sia, who explained that the
 license was comparable to the hypothetical negotiation be-
 cause the AppliedBio license covered reagents that would
 enable a researcher to perform PCR in an improved man-
 ner, and the patents-in-suit also deal with performing im-
 proved PCR reactions using droplet technology. The jury
 also heard the testimony of Bio-Rad’s corporate witness,
 Ms. Tumolo, who compared the two technologies and testi-
 fied that the $0.12 per-unit royalty for the improved rea-
 gents in the AppliedBio agreement would translate to a
 much higher royalty rate per reaction in the hypothetical
 negotiation because one would use more quantities of the
 reagents in microfluidic systems. Accordingly, the evi-
 dence presented at trial regarding this agreement also sup-
 ports the 15% reasonable royalty rate. 5



     5   Having concluded that substantial evidence sup-
 ports the jury’s verdict of a 15% royalty rate, we need not
 address the parties’ arguments regarding the
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     Finally, we note that 10X’s reliance on our decisions in
 LaserDynamics and ResQNet in support of its arguments
 is misplaced. In LaserDynamics, for example, the royalty
 rate proposed by the expert was “untethered from the pa-
 tented technology at issue and the many licenses thereto.”
 694 F.3d at 81. Likewise, ResQNet involved a royalty
 award based entirely on inapposite licenses. 594 F.3d at
 872. In both cases, the expert used licenses that served no
 purpose other than “to increase the reasonable royalty rate
 above rates more clearly linked to the economic demand for
 the claimed technology.” LaserDynamics, 694 F.3d at 80.
 Here, by contrast, Mr. Malackowski evaluated the various
 licenses at issue and applied the Georgia-Pacific factors.
 He testified that it made sense to adopt a 15% reasonable
 royalty rate where the parties to the hypothetical negotia-
 tion are direct competitors. He also provided specific rea-
 sons why the University of Chicago/RainDance license was
 not comparable. Accordingly, we conclude that substantial
 evidence supports Mr. Malackowski’s reasonable royalty
 opinions and the jury’s verdict.
            B. 10X’s Apportionment Argument
                     Is Without Merit
     “When the accused technology does not make up the
 whole of the accused product, apportionment is required.
 The ultimate combination of royalty base and royalty rate
 must reflect the value attributable to the infringing fea-
 tures of the product, and no more.” Elbit Sys. Land & C4I
 Ltd. v. Hughes Network Sys., LLC, 927 F.3d 1292, 1301
 (Fed. Cir. 2019) (internal quotations and alterations omit-
 ted). “[A] reasonable royalty analysis necessarily involves




 comparability of the Applera/Bio-Rad license. As discussed
 above, we do not think the district court abused its discre-
 tion in allowing the jury to hear testimony regarding that
 license.
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 an element of approximation and uncertainty.” Lucent
 Techs., 580 F.3d at 1336 (internal quotations omitted).
     10X challenges Mr. Malackowski’s testimony because
 of his alleged failure to apportion damages between the pa-
 tented and unpatented features of the accused products. In
 10X’s view, Mr. Malackowski claimed that his 15% royalty
 rate was already apportioned in the comparable licenses,
 but failed to provide any numerical value to support his
 analysis. 10X also argues that none of the other witnesses
 provided any testimony that could fill the gaps as to the
 technical contributions of any of the patents. We disagree.
     As Bio-Rad correctly points out, there is no blanket rule
 of quantitative apportionment in every comparable license
 case. In Elbit Systems Land & C4I Ltd. v. Hughes Network
 Systems, LLC, for example, we accepted “built in appor-
 tionment” for a comparable license agreement. 927 F.3d at
 1301 (internal quotations omitted). 10X argues that Elbit
 is distinguishable because the license at issue in that case
 was the “closest” comparator and the expert in Elbit actu-
 ally made a quantitative adjustment to the comparator li-
 cense. Appellant’s Reply Br. 31. But this argument rests
 primarily on the faulty assumption that the Chi-
 cago/RainDance license (with the 1–3% royalty rate) is the
 most comparable license in this case. As discussed above,
 the jury was free to accept Bio-Rad’s evidence that this li-
 cense was not comparable.
     Here, Mr. Malackowski concluded that no quantitative
 adjustment of the royalty rate in the three agreements was
 required. He explained that his methodology involved
 looking at comparable license agreements between compet-
 itors for similar technologies and assessing whether the
 importance of that technology to the particular license was
 similar to the hypothetical negotiation. He also acknowl-
 edged that he relied on the reports, testimony, and conclu-
 sions of other witnesses to understand that the licenses
 were technologically comparable, and that the proportion
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 34             BIO-RAD LABORATORIES, INC.   v. 10X GENOMICS INC.



 of licensed/unlicensed features was comparable to the pre-
 sent case. Thus, under Mr. Malackowski’s reasoning, no
 adjustment of the 15% royalty rate in the comparable li-
 censes was required. His analysis could reasonably be
 found to incorporate the required apportionment. Our case
 law does not require more. See, e.g., VirnetX, Inc. v. Cisco
 Sys., Inc., 767 F.3d 1308, 1328 (Fed. Cir. 2014) (“[W]e note
 that we have never required absolute precision in [applying
 the principles of apportionment]; on the contrary, it is well-
 understood that this process may involve some degree of
 approximation and uncertainty.”); see also Ericsson, Inc.,
 773 F.3d at 1227 (recognizing that, even though “[p]rior li-
 censes . . . are almost never perfectly analogous to the in-
 fringement action,” if accompanied by testimony
 accounting for the distinguishing facts, prior licenses may
 help the jury decide an appropriate royalty award).
     This is not a case in which an unsupported conclusory
 opinion leaves the jury with nothing but speculation. We
 thus agree with the district court that Mr. Malackowski’s
 testimony was properly admitted. We therefore affirm the
 damages award.
                         D. INJUNCTION
     “According to well-established principles of equity, a
 plaintiff seeking a permanent injunction must satisfy a
 four-factor test before a court may grant such relief.” eBay
 Inc. v. MercExchange, LLC, 547 U.S. 388, 391 (2006). “A
 plaintiff must demonstrate: (1) that it has suffered an ir-
 reparable injury; (2) that remedies available at law, such
 as monetary damages, are inadequate to compensate for
 that injury; (3) that, considering the balance of hardships
 between the plaintiff and defendant, a remedy in equity is
 warranted; and (4) that the public interest would not be
 disserved by a permanent injunction.” Id. The district
 court’s grant of an injunction is reviewed for abuse of dis-
 cretion. eBay Inc., 547 U.S. at 391. We review the district
 court’s conclusion as to each eBay factor for abuse of
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 discretion and its underlying factual findings for clear er-
 ror. i4i Ltd. P’ship v. Microsoft Corp., 598 F.3d 831, 861
 (Fed. Cir. 2010), aff’d, 564 U.S. 91 (2011). Only the first,
 third, and fourth factors are at issue on appeal.
     The district court found that Bio-Rad satisfied all four
 eBay factors. On appeal, 10X argues that Bio-Rad failed to
 justify its request for a permanent injunction because it did
 not show irreparable harm, and because neither the bal-
 ance of hardships nor the public interest support an injunc-
 tion. We address these arguments below.
                    1. Irreparable Harm
     To prove irreparable injury, a patentee must show
 “that absent an injunction, it will suffer irreparable harm,
 and . . . that a sufficiently strong causal nexus relates the
 alleged harm to the alleged infringement.” Apple Inc. v.
 Samsung Elecs. Co., 695 F.3d 1370, 1374 (Fed. Cir. 2012).
     The district court found that 10X and Bio-Rad are di-
 rect competitors, and Bio-Rad would suffer irreparable
 competitive harm absent an injunction. 10X argues, how-
 ever, that it was improper for the court to enjoin sales in
 the name of competition because 10X and Bio-Rad do not
 directly compete. 10X states that it is undisputed that Bio-
 Rad did not even claim to compete with four out of five of
 10X’s product lines. According to 10X, only one product
 line, 10X’s Single Cell 3’, potentially competes with Bio-
 Rad ddSEQ product. Even as to this product, however, 10X
 argues that there is no irreparable harm because Bio-Rad’s
 product is inferior, faces competition from at least ten other
 competitors, and Bio-Rad could not prove that it lost a sin-
 gle sale to 10X. Thus, in 10X’s view an injunction is un-
 likely to help Bio-Rad’s competitive position. Appellant’s
 Br. 73–74.
     Bio-Rad responds that 10X admitted that the Single
 Cell 3’ product, which accounts for over 80% of 10X’s reve-
 nue, competes directly with Bio-Rad’s ddSEQ product. Bio-
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 36             BIO-RAD LABORATORIES, INC.   v. 10X GENOMICS INC.



 Rad also contends that 10X’s allegations of commercially
 and technically superior products support Bio-Rad’s claim
 of irreparable harm. Bio-Rad explains that it was forced to
 market early in view of the head start 10X received from
 its willful infringement. Bio-Rad also explains that it “fell
 behind 10X in the droplet field because the 10X founders
 left Bio-Rad to start 10X with much of Bio-Rad’s droplet
 expertise and hired away even more droplet-experienced
 technologists.” Appellees’ Br. 58–59. In Bio-Rad’s view,
 the first mover advantage allowed 10X to capture many
 “sticky” customer relationships and secure a competitive
 lead. Finally, Bio-Rad contends that 10X’s argument that
 Bio-Rad did not lose any customers is inconsistent with the
 fact that the two company’s products are sold “head to
 head.” Id. at 57–61.
     It is undeniable that Bio-Rad has suffered harm from
 10X’s first mover advantage and “sticky” customer rela-
 tionships. The district court found that Bio-Rad is being
 forced to compete with 10X’s products that incorporate the
 infringing technology. Based on its willful infringement—
 a finding 10X does not challenge on appeal—10X has es-
 tablished a strong market lead over Bio-Rad. The court
 also found that, based on 10X’s first mover advantage, Bio-
 Rad had to increase its marketing costs. Money damages
 will not be able to compensate Bio-Rad for the harms stem-
 ming from 10X’s first mover advantage.
                    2. Balance of Hardships
     In considering the balance of hardships, courts may
 consider the “parties’ sizes, products, and revenue sources.”
 i4i Ltd. P’ship, 598 F.3d at 862. “[E]xpenses . . . incurred
 in creating the infringing products” and “the conse-
 quences . . . of its infringement, such as the cost of rede-
 signing the infringing products” are “irrelevant.” Id. at
 863.
    The district court found that the balance of hardships
 weighed in favor of injunctive relief or, at minimum, was
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 neutral. The court found that Bio-Rad’s hardship stems
 from the fact that it significantly invested in its droplet
 business. And the court found that 10X’s hardship is miti-
 gated because it could sell its new, non-infringing products.
 On appeal, 10X argues that the district court failed to con-
 sider that it does not have a new design for two of its prod-
 uct lines. 10X explains that it is a much smaller company
 than Bio-Rad and its entire business depends on the en-
 joined products.
      Bio-Rad contends that 10X’s argument that 10X does
 not have a new design for two of its product lines (Linked-
 Reads and CNV) is undermined by the fact that 10X’s
 Linked-Reads and CNV products are trivial to its econom-
 ics. Bio-Rad also points out that the district court gave 10X
 a fair amount of time to design around the patents-in-suit.
 We do not agree.
     We acknowledge that Bio-Rad, although a much larger
 company, will suffer considerable hardship absent an in-
 junction because it has invested almost half a billion dol-
 lars to develop its products, including acquisitions and tens
 of millions of dollars a year on research and development.
 But at the same time, 10X, a much smaller company, de-
 pends entirely on the sales of the enjoined products for its
 revenue. Although the district court concluded that the
 hardship to 10X is mitigated because it can sell its non-in-
 fringing alternatives, the district court failed to consider
 the lack of non-infringing products for two out of the five
 product lines. In the absence of non-infringing alternatives
 for the Linked-Reads and CNV products, we conclude it
 was an abuse of discretion for the district court to enjoin
 sales of these two product lines.
                      3. Public Interest
    “[T]he touchstone of the public interest factor is
 whether an injunction, both in scope and effect, strikes a
 workable balance between protecting the patentee’s rights
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 38             BIO-RAD LABORATORIES, INC.   v. 10X GENOMICS INC.



 and protecting the public from the injunction’s adverse ef-
 fects.” i4i, 598 F.3d at 863.
     The district court carved out an exception for infringing
 products that were sold or in use before the effective date
 of the injunction (the “Historical Installed Base”). For
 these products, 10X can also continue to supply consuma-
 bles, and support, service, repair, and replace them under
 warranty. This exception is conditional on 10X paying a
 15% royalty on the net revenue 10X receives from the per-
 mitted Historical Installed Base sales until the expiration
 of the patents-in-suit. Despite this carve-out, 10X argues
 that the undisputed evidence demonstrates that scientists
 need 10X products to do important research, which weighs
 against injunctive relief. Appellant’s Br. 78–79.
      We see no abuse of discretion here. Under the district
 court’s injunction, 10X’s existing customers are not en-
 joined from using their installed systems, as long as 10X
 pays royalties. Thus, there is no basis for 10X’s argument
 that scientists will lose their years of research or be finan-
 cially precluded from working on existing projects. On-go-
 ing research projects will not be affected, and the scientist
 statements submitted by the amicus all presuppose that
 they will be required to give up their existing equipment.
 None of the letter writers, moreover, seem to be aware of
 10X’s replacement non-infringing design that supposedly
 works just as well as the Historically Installed Base.
     The district court carefully crafted an injunction that
 allows existing 10X customers to continue their important
 research but attempts to mitigate the harm to Bio-Rad
 from 10X’s first mover advantage. The court also provided
 10X with an opportunity to design non-infringing alterna-
 tives, which 10X has done for all but two infringing product
 lines. In these circumstances, we conclude that the district
 court did not abuse its discretion in granting Bio-Rad an
 injunction, except as to the Linked-Reads and CNV product
 lines. Accordingly, we vacate the injunction as to those two
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 product lines only, but conclude that the injunction should
 remain in place as to the other enjoined product lines.
                        CONCLUSION
      We have considered the parties’ remaining arguments
 and find them unpersuasive. For the foregoing reasons, we
 affirm-in-part, reverse-in-part, and vacate-in-part the dis-
 trict court’s decision denying 10X’s motion for JMOL. We
 vacate the district court’s injunction only with respect to
 10X’s Linked-Reads and CNV product lines.
     AFFIRMED-IN-PART, REVERSED-IN-PART,
       VACATED-IN-PART, AND REMANDED
                              COSTS
     No costs.
