The Plica Semilunaris

The plica semilunaris is a small fold of conjunctiva that steers debris out of the eye and carries immune tissue, so it is not useless. Its match with the third eyelid of other mammals is well supported. It is reduced against lemurs and lorises, but whether that primate pattern reflects ancestry or need has not been tested.

By AI (Claude Opus 5.5, reasoning effort 40); strengthening review by Claude Opus 5.5, thinking level unavailable Draft:  September 2026

"There can be no doubt that this is brought about by the intervention of the plica semilunaris, which is not so vestigial a structure as the descriptions of it might imply" — E. Philip Stibbe, Journal of Anatomy, 19281

Summary

The parent article, Vestigial Structures, separates two claims.  The old claim says a vestige is useless.  The live claim says a vestige is a reduced copy of a fuller structure in related animals, and it has three steps: homology, reduction, and a distribution across species that fits a family tree built from other characters.  This article runs that test on the small pink fold at the inner corner of the eye.

  • What it is. A crescent of conjunctiva about 2 mm wide at the inner corner of the eye.2 Gray's Anatomy of 1918 reports that "Müller found smooth muscular fibers in this fold; in some of the domesticated animals it contains a thin plate of cartilage."3 In 2004 G. Arends and U. Schramm sectioned it in eleven fetuses, two newborns and an old man, and found goblet cells, blood vessels and dense immune cells, but no cartilage and no muscle outside the vessel walls.44
  • What it does. In 1928 the Liverpool anatomist E. Philip Stibbe traced dust to "a little sticky mass at the inner canthus" and watched the fold catch a loose eyelash and pass it to the skin.11 Nobody has measured this, but the useless claim does not survive it.
  • The third eyelid elsewhere. Birds draw a clear membrane across the eye with two muscles supplied by the sixth cranial nerve.1 Mammals lack them; their third eyelid, stiffened by cartilage in horses, cattle and cats, "slips over the eye only passively when the globe is slightly or markedly retracted."5
  • Primates. Tatsuyoshi Arao and Edwin Perkins examined 22 non-human primate species in 1968. Twenty-one had "a well developed nictitating membrane," covering from the whole eye in the potto to a tenth of it in the gorilla; only the chimpanzee's was vestigial. Nearly all prosimians and three New World monkeys had cartilage in it.6
  • Two claims. Darwin called the fold "a mere rudiment" of the third eyelid, a homology claim.7 S. R. Scadding in 1981 answered only the function claim; Bruce Naylor replied in 1982 that the fold is "powerful suggestive evidence" of descent.89
  • The creationist replies. Creation Ministries International, Answers in Genesis and the Institute for Creation Research answer the useless claim, rightly on the main point.101112 Answers in Genesis says the fold "is thought to be homologous" and does not dispute it.13 None discusses the primate data.
  • The test. Homology is strong. Reduction against lemurs and lorises is real. The third step is weak: the primate pattern follows the family tree, but it also follows how much each animal needs a third eyelid, and no study has separated the two.

Verdict: the plica semilunaris is not useless, and creationists are right about that.  It is very probably the human form of the mammalian third eyelid, reduced in structure, and Answers in Genesis does not contest that.  Whether the reduction records ancestry or a design fitted to a face with a mobile upper lid and no eyeball-retracting muscle cannot be decided from the published data.  The fold is weak evidence for either side.

Part 1: What the fold is and what it does

Anatomy

The conjunctiva is the thin, moist membrane that lines the eyelids and covers the white of the eye.  Gray's Anatomy of 1918 says that "at the medial angle it forms a semilunar fold, the plica semilunaris."3  On the nose side of the fold sits the caruncle, a pink mound of modified skin that fills the "lacus lacrimalis," the small pool where tears gather before they drain through two pinhole openings, the puncta, in the lid margins.3  The fold's free edge is curved, with its hollow side facing the cornea.

It is small.  Wiedersheim wrote in 1895 that it is broader in the newborn "and during the early years of life," and later "does not exceed 1½ to 2 mm. in breadth."2  It is deeper than it is wide.  Stibbe called it "a very respectable fold, with an underlying conjunctival fornix often a quarter of an inch in depth."1

Tissue

The fullest modern histology found is Arends and Schramm's 2004 study of fourteen people, from fetuses to an old man, read here in abstract because the full paper is paywalled.  It reports five findings.

  1. Early size. Early in development the fold "covered a bigger part of the orbit and later did not keep up with the growth of the eyeball and the lids."4
  2. Gland buds. They found "poorly differentiated buds of the surface epithelium which can be classified as rudiments of the nictitating or Harderian gland," plus serous glands that "could be beginnings of Krause's glands."4 The Harderian gland serves the third eyelid in other animals.
  3. Mucus. The side facing the lids had more cell layers "and more goblet cells than its bulbar side."4 Goblet cells secrete mucus.
  4. No cartilage, no muscle. The connective tissue held "no elastic or reticular fibres or cartilaginous structures." Some blood vessels had a thin coat of muscle cells, but otherwise "Muscle cells could not be detected."4
  5. Immune cells. "A very dense concentration of leukocytes" was present even before birth, some arranged as follicles.4

The fourth finding conflicts with two older reports.  Gray repeats Müller's report of smooth muscle fibres in the fold.3  Wiedersheim says the fold "may still enclose (more frequently in Negroes than in Caucasians) a cartilaginous support," and that Giacomini found this cartilage in twelve of sixteen Black people he examined.22  Stibbe looked for it and did not find it: "The writer has as yet not found any cartilage in a number of sections of the plica semilunaris."1  Arends and Schramm's fourteen specimens had none either.  No modern count of how often a human fold contains cartilage, in any population, was found.

Function

Stibbe's 1928 paper is the only direct observation of the fold at work found for this article.  He began with a dust storm: "one's eyes rapidly fill with dust, and as rapidly get rid of it.  Where does the dust go?  It will be found collected together into a little sticky mass at the inner canthus".1  Tears and blinking sweep particles to the inner corner, past the puncta, which are "about one-tenth of a millimetre in diameter."  The fold bridges the pocket of conjunctiva there, "So the tears and foreign matter are diverted forwards across the caruncular region," where the caruncle's fine hairs and greasy secretion hold them on the skin.1

He also watched an eyelash in a patient's eye, which "slipped under the plica semilunaris; it was brought out from this position by an outward sweep of the eye abolishing the deep internal fornix; and after several trials it was finally picked up by the plica, and transferred to the skin at the inner canthus."1  The same sentence bears on a second job sometimes claimed for the fold: when the eye turns outward the pocket under it flattens, so the fold is spare conjunctiva that lets the eye rotate freely.  That is an inference from Stibbe's description, not a measurement.

Arends and Schramm add the immune and mucus roles: the fold "plays an important role as a specialized organ in human eye protection."4  The immune cells they saw belong to a known system.  In 2000 Nadja and Erich Knop examined 53 whole conjunctival sacs from donor eyes.  Every one carried a thin layer of lymphocytes and mostly IgA-making plasma cells, about three fifths also had organized follicles, and together these hold "all components necessary for a complete immune response."14  That makes the fold's leukocytes working tissue, not debris of an old organ.  It also limits the claim:  the Knops found this tissue "mainly" in the conjunctiva lining the lids, so the plica is one part of a sheet that covers the whole eye surface, not its immune centre.14  None of this is quantified for the fold itself.  No study was found that measured debris clearance, tear drainage or eye rotation with and without a working fold.  The evidence is enough to end the claim that the fold does nothing, not to say how much it does.

Part 2: The third eyelid in other animals

Birds and reptiles

Gordon Walls's 1942 survey, The Vertebrate Eye and Its Adaptive Radiation, describes the third eyelid, or nictitating membrane, as "a fold of the conjunctiva at the inner or nasal corner of the lid opening" that, "Being transparent, it can sweep the cornea from the nasal to the temporal side, to clean and moisten it, without shutting out the light."5  In birds the inner surface is lined with cells "built like unicellular feathers, which improve its cleansing action."5

Birds work it with muscles.  Stibbe dissected the ostrich and traced the pyramidalis, whose tendon wraps round the eyeball to the membrane, and the quadratus, which holds that tendon off the optic nerve.  "The nerve supply of the pyramidalis and of the quadratus is from the sixth cranial nerve".1

Mammals

Mammals have a third eyelid of a different build.  Walls: "Where it is well developed, it usually has a cartilaginous tarsus, but it never has a special musculature behind the globe as in lower vertebrates.  Hence, it slips over the eye only passively when the globe is slightly or markedly retracted."5  Septimus Sisson's description of the horse shows the mechanism.  The third eyelid is "a semilunar fold of the conjunctiva, known as the membrana nictitans, which covers and partly encloses a curved plate of hyaline cartilage," with "Numerous minute lymph nodules" and a gland.  When the eyeball is pulled back, it presses on the fat around the cartilage and the membrane is pushed out over the eye until about an inch of it shows.15  Sisson's words for the horse, a semilunar fold of the conjunctiva at the inner corner, are Gray's words for the human.

The cat adds smooth muscle.  J. W. Thompson wrote in 1961 that the cat's membrane "contains two thin sheets of smooth muscle" inserted "into adjacent sides of a T-shaped piece of cartilage," and these muscles pull it back.1616  Müller's smooth muscle in the human fold, if it is there, would be the counterpart.  In cattle, sheep, rabbits and cats Stibbe found no muscle that moves the membrane, and in serial sections "at the most a few scattered bundles of unstriped muscle fibres."1

Across mammals the third eyelid is patchy.  All carnivores but the skunk have one, the cat's haw reaching right across the cornea, and all ungulates have one, "usually with no apparent usefulness," while Walls lists it as "vestigial in rodents and others of the lower orders such as the insectivores, primates, and 'edentates'."555

Primates

Walls's "vestigial in ... primates" does not match the one survey that looked closely.  Arao and Perkins's 1968 paper in The Anatomical Record, of which only the abstract could be obtained, found "a well developed nictitating membrane" in 21 of 22 non-human species.  It was "vestigial only in Pan satyrus/troglodytes," the chimpanzee, and ranged from a tenth of the eye in the gorilla to the whole eye in the potto and the angwantibo.  Cartilage was present in every prosimian studied except the thick-tailed galago, and in three New World monkeys: the red uakari, the white-fronted capuchin and the squirrel monkey.6  "Muscle fibers are present only in the nictitating membranes of Lemur monoz and Callithrix aurita."6

So among primates the fullest third eyelids belong to the lorises and lemurs, the monkeys have smaller ones, and the human fold sits with the great apes.  Its lack of muscle is normal for a primate.

The membrane does not only shrink along that series.  It changes build.  Arao and Perkins found that "The outer surface of the nictitans of all species of anthropoids has mucous glands; that of the prosimians has few of them."6  The human fold fits the anthropoid side:  goblet cells crowd its lid-facing surface, and Arends and Schramm found in it "a new kind of plica gland," with ducts distinct from the mucous glands.44  In monkeys, apes and humans the smaller membrane is the more glandular one.

The gland follows a similar course.  Walls wrote that neither the special muscles nor "the special gland (Harder's) of the third eyelid are present, even as vestiges, in man."5  Susan Rehorek and Timothy Smith showed in 2006 that this is wrong for the unborn: the Harderian gland is "largely absent in the adult human," but "it is present in the fetal and neonatal stages."  In the newborns of twelve other primate species its size ranged "from large to absent (tarsiers), with no apparent phylogenetic trends."1717

Is the bird's membrane the same organ?

Stibbe doubted it.  He concluded "That structurally and functionally the plica of man and Mammals differs entirely from the nictitating membrane of Birds," and left the question of common origin open until someone studied its development.1  Walls treats them as one organ whose muscles the early mammals lost.5  Stibbe did not doubt that the human fold belongs with the mammalian one, and the human case rests on mammals and above all primates.  The bird question does not decide anything here.

Part 3: Two claims about one fold

Darwin's 1871 statement is about homology and reduction, not use.  He traces the third eyelid from birds, where it is "of much functional importance," through reptiles, sharks, monotremes, marsupials and the walrus, to man, where "it exists, as is admitted by all anatomists, as a mere rudiment, called the semilunar fold."7  He never calls it useless.

Wiedersheim in 1895 made both claims.  His text is about homology: the fold "corresponds with the third eyelid, the so-called nictitating membrane, of the lower animals," and in man and the apes "this third eyelid has undergone great degeneration."  He also gave a reason: in those animals it cleans the eye, "the upper lid [which in Man performs that function] being immovable."2  His conspectus then put the fold in section B, organs that have become "wholly or in part functionless."2

Scadding's 1981 reply took up only the second claim.  The fold "is simply that portion of the conjunctiva at the medial corner of the eye and as such aids in the cleansing and lubrication of the eye ball," and "The absence of a nictitating membrane in humans is of no significance to a discussion of the function of the semilunar fold."8  The parent article calls that second sentence "the wrong-claim error in miniature."  The charge holds, with one qualification.  As a sentence about function it is true, and Scadding did not pretend to refute the homology; he set it aside.  But his paper concluded that vestigial organs "provide no special evidence for the theory of evolution,"8 and for the plica that needs the homology answered, since Darwin's version rested on homology alone.

Naylor saw this.  His 1982 reply: "the fact remains that the coccyx, ear muscles, and semilunar fold of the eye provide powerful suggestive evidence for the evolution of man from "lower" primates."9  Jerry Coyne's 2009 definition, that a trait "can be vestigial and functional at the same time,"18 is Naylor's.  Coyne's book does not use the plica at all.  Searches of its text for "plica," "semilunar," "nictitating" and "third eyelid" returned nothing.

Part 4: The creationist replies

In their own words first.

  • Robert Franks, Creation magazine, 1988. "Some evolutionists feel that it is a remnant of the third eyelid of a lower form, such as the third eyelid in birds and reptiles. But in man this tissue has several essential functions. If you did not have the tissue for support at that location, the eyeball would sink. This would cause double vision. The tissue not only supports the eyeball, but the tearduct as well." It "also serves to collect foreign matter." "It was designed. And it does not represent the cleverly designed third eyelid of the bird".1010
  • Jerry Bergman, 2000. A figure lists the plica among "some of the 180 structures that were considered vestigial" and says "most of these structures have at least one function."19 Bergman and George Howe's 1990 book on vestigial organs could not be obtained.
  • Elizabeth Mitchell, Answers in Genesis, 2011. "While this fold of protective eye covering is thought to be homologous with the more complete protective covering in some animals, it is far from useless. Secretions from this "#1 useless body part" mop debris from your eyes." Her explanation of such homologies: "our common Designer used well-engineered designs to serve various functions."1313
  • Lita Sanders, Creation Ministries International, 2011. "The plica semilunaris actually has a critical function for the health of the eye. It secretes the sticky substance that hardens into the crust we sometimes find around our eyes when we wake up."11
  • Randy Guliuzza, Institute for Creation Research, 2025. The plica "or 'third eyelid'" is one of the obscure parts in what he calls the "vestigial organs scam," whose recipe includes declaring a part vestigial "since no one knows of a useful function."12

On function these writers are right in substance: the debris-collecting role is Stibbe's, and the mucus and immune tissue are in Arends and Schramm.  Two claims go too far.  Franks's statement that without the fold "the eyeball would sink," causing double vision, carries no source.  None of the anatomical sources read for this article gives a 2 mm fold of conjunctiva any role in holding up the eyeball.  Sanders credits the fold with the morning crust, but Gray names the caruncle as "the source of the whitish secretion which constantly collects in this region."3  The fold's goblet cells add mucus, so the claim is partly right.

On homology, the replies say little.  Mitchell leaves it standing.  Franks contrasts the fold with the bird's membrane, which Stibbe also did, but the human claim rests on mammals and primates, and no reply mentions them.  Guliuzza treats a vestige claim as resting on "no one knows of a useful function," which is the 1895 claim, not Coyne's.  The reply from this literature that could meet the modern claim is Sarfati and Batten's: a lost or reduced organ "would prove devolution not evolution.  The creation model allows for deterioration of a perfect creation."20  That allows reduction within a created kind.  It cannot cover this case, because the comparison runs from humans to lemurs and lorises, and young-earth creationists hold that humans share a created kind with no other primate.  The design reply has to say that the Creator gave each primate the fold its eyes need, and nobody has written that argument up with data.

Part 5: The three-step test

Step 1: homology is strong

The fold sits where the third eyelid sits in every mammal that has one, and anatomists describe the horse's and the human's in the same words.153  Some human folds have been reported to contain cartilage, the stiffening plate of the mammalian third eyelid.2  The fetal fold holds buds that Arends and Schramm read as rudiments of the third eyelid's gland, and the Harderian gland itself is present in the human fetus and newborn.417  Like the horse's, it carries immune tissue.15  Arends and Schramm, who were arguing for its function, still wrote that "Its origin from the third eyelid of the mammals could be recognized to a certain extent."4  Answers in Genesis does not contest the homology.  The bird comparison is doubtful, but it is not needed.

Step 2: reduction is real, with limits

Against a loris or a lemur, the human fold is reduced.  It covers none of the cornea, usually has no cartilage, and the adult has largely lost the gland.  Two of Darwin's diagnostics apply.  The fold is relatively larger early in fetal life, and it varies in size and in whether it holds cartilage.24

Two limits apply.  First, the human fold is not exceptional among its nearest relatives; the chimpanzee's is also vestigial and the gorilla's covers a tenth of the eye.6  Second, what is reduced is structure and reach, not clearly the job.  Stibbe's point was that the mammalian third eyelid, even in the calf, does not sweep the eye like a bird's; it intercepts debris, and the human fold does that too.1  The gland data point the same way.  The anthropoid membranes, human fold included, are smaller and more glandular than the prosimian ones, so the change is from a sliding shield toward a secreting, debris-catching fold, not plain loss.64  This answers the picture of the fold as a decaying leftover.  It does not answer the ancestry claim.  The glandular build is shared by monkeys, apes and humans, so it also fits the tree, and a skeptic can read it as a new role taken on by an inherited part, which descent allows.

A 2017 case report from India describes a nine-year-old girl born with a membrane 20 mm by 18 mm spreading from the inner corner over a third of her left cornea, which the authors call a persistent nictitating membrane, the second human case they found.2121  But the removed tissue was plain conjunctiva: "No elastic, reticular fibers, cartilaginous structures, or muscle cells could be found".21  That looks like an overgrown fetal fold, not the return of an ancestral organ, and it adds little either way.

Step 3: the ancestral explanation is the weak step

Lorises and lemurs have the fullest third eyelids, nearly all with cartilage; some New World monkeys keep cartilage; the gorilla's is small, the chimpanzee's vestigial, and the human's a narrow fold.6  That fits a tree on which the membrane shrank along the line to the apes, which is what descent predicts.

Three things keep it from being strong.

  1. Need predicts much of the same pattern. A mammal's third eyelid slides across the eye only when the eyeball is pulled back.5 Humans lack the muscle that does the pulling. Wiedersheim linked the fold's reduction to "the absence of a retractor bulbi muscle," and Walls wrote that "In man, it is the heavy bony rim of the orbit, particularly the ridge bearing the eyebrow, which makes a retractor bulbi unnecessary."25 Wiedersheim also noted that the human upper lid does the cleaning. A large third eyelid on such a face would have nothing to push it out, so a design fitted to need also predicts a small fold. Walls himself explained the horse's large third eyelid by grazing in deep grass.5
  2. Across mammals the organ does not shrink in one direction. Walls found it "most rudimented in the lower mammals" and back to "greatest usefulness in some of the higher ones."5 On a tree that means repeated loss or regrowth, which weakens any one-way story. In the primate newborns Rehorek and Smith examined, the Harderian gland's size showed "no apparent phylogenetic trends."17
  3. The key data are thin. Arao and Perkins's species table was not available, and no later survey was found.

What would decide it is a comparative study that codes third-eyelid size, cartilage and gland across many primates and tests, on a phylogeny, whether they follow ancestry after activity pattern, orbit shape and the retractor bulbi are accounted for.  Descent predicts a signal of ancestry left over; a design fitted only to need predicts none.  A modern count of cartilage in human folds, and a study of the genes that build the third eyelid, would also help.  None was found.

Objections and responses

  • "The fold works, so it is not a vestige." That answers the 1895 claim. Coyne's definition allows a vestige to work.18 Function does not touch homology or ancestry.
  • "Similar parts show a common Designer." Common design explains similarity. It does not, without more, explain why the human fold is relatively larger before birth or holds buds of a gland the adult lacks. No design account of those has been offered.
  • "A larger fetal fold is Haeckel's discredited recapitulation." No. It is Darwin's diagnostic that a reduced organ is often relatively larger in the embryo, not a claim that embryos replay adult ancestors. It is still weak alone, since organs grow at different rates.
  • "The chimpanzee's third eyelid is vestigial and the gorilla's is not, so the pattern breaks the tree." Further loss on one branch is allowed by descent. But the pattern is then not a simple trend, which is part of why step 3 is weak.
  • "A girl was born with a full third eyelid, an atavism." Her membrane had no cartilage, muscle or gland.21 It is weak evidence either way.
  • "Stibbe showed that mammals have no true nictitating membrane, so the homology fails." Stibbe separated mammals from birds, not humans from mammals.1
  • "The design-by-need reading is invented after the fact." Fair. It fits the anatomy, but no creationist has tested it, so it is an alternative, not a result. The descent reading has the matching gap: it has not been shown to beat need on the primate data.

Issues and unanswered questions

  • No measured function. Debris interception rests on Stibbe's observations, and the immune and mucus roles on tissue. How much of the eye surface's immune tissue sits in the fold, against the lid lining where the Knops found most of it, has not been counted.
  • Primary data unread. Only the abstracts of Arao and Perkins 1968 and Arends and Schramm 2004 were obtained. The species list, and whether Arao and Perkins compared humans, are unknown.
  • Cartilage and muscle in humans. The cartilage reports are nineteenth-century counts framed by race, known here second-hand; Stibbe and Arends and Schramm found none. Müller's smooth muscle was not found by Arends and Schramm. Both questions are open.
  • No phylogenetic test. No study tests whether primate third-eyelid size tracks ancestry or ecology.
  • Creationist gaps. Bergman and Howe 1990 was not obtained. Franks's claim that the eyeball would sink without the fold has no support in any source read here. No creationist source discusses the primate data or offers a need-based account with numbers.

Verdict

The useless claim is wrong.  The fold steers debris to the inner corner and carries mucus-secreting and immune tissue, and creationists were right to say so.  The homology claim is strong.  The fold matches the third eyelid of other mammals in position, tissue, occasional cartilage and fetal gland, and Answers in Genesis does not dispute it.  Reduction relative to lemurs and lorises is real.  What the fold cannot do is carry the ancestral step on its own: the primate pattern follows the tree and follows need, and nobody has tested which fits better.  The plica semilunaris is weak evidence for either side.

Sources


  1. 1.

    Stibbe, E. Philip.  “A comparative study of the nictitating membrane of birds and mammals.”  Journal of Anatomy 62(Pt 2):159–176.  1928.  A comparison by E. Philip Stibbe, an anatomist at the University of Liverpool, of the third eyelid in birds, which has its own muscles, with the same membrane in calves, sheep, rabbits and cats, which has no special muscle.  He also describes the human fold catching dust and foreign bodies, and doubts that it is the same structure as a bird's membrane.  The article uses him for the dust-catching role and for the muscle difference between birds and mammals.

    1. 1a.

      Pages 169–170:  “If one goes out in a dust storm, one's eyes rapidly fill with dust, and as rapidly get rid of it.  Where does the dust go?  It will be found collected together into a little sticky mass at the inner canthus; and that mass is situated on skin, in the angle internal to the caruncula lachrymalis, in which situation it causes no irritation and does no damage.  There can be no doubt that this is brought about by the intervention of the plica semilunaris, which is not so vestigial a structure as the descriptions of it might imply; it is in fact a very respectable fold, with an underlying conjunctival fornix often a quarter of an inch in depth.” ↩ ↩ ↩ ↩

    2. 1b.

      Page 170:  “An eyelash, for example, has been observed in a patient's eye; it was carried inwards by the tears and the action of the orbicularis, but slipped under the plica semilunaris; it was brought out from this position by an outward sweep of the eye abolishing the deep internal fornix; and after several trials it was finally picked up by the plica, and transferred to the skin at the inner canthus.” ↩ ↩

    3. 1c.

      Page 170:  “(a) that the orifices of the canaliculi are very minute—about one-tenth of a millimetre in diameter”.  “Now the internal conjunctival fornix is bridged across by the plica semilunaris, which in man has a very thin free concave edge, like the lunula of a watch-pocket valve of the heart.  So the tears and foreign matter are diverted forwards across the caruncular region.” ↩

    4. 1d.

      Page 164:  “The nerve supply of the pyramidalis and of the quadratus is from the sixth cranial nerve, which supplies the external rectus, and then turns inwards and backwards under the quadratus to end in the pyramidalis (fig. 2).” ↩ ↩

    5. 1e.

      Pages 164–165:  “The writer has made numbers of dissections of the orbits of calves, sheep, rabbits and cats, all of which have a well-marked "nictitating membrane," and has failed to find any specialised apparatus for moving the membrane.  The examination of serial sections across the fold has revealed at the most a few scattered bundles of unstriped muscle fibres.” ↩

    6. 1f.

      Page 175:  “In conclusion, it has been assumed that the plica semilunaris of man is the same as the plica of Mammals; cartilage has been found in the plica of negroes, and occasionally in Europeans.  The writer has as yet not found any cartilage in a number of sections of the plica semilunaris.” ↩

    7. 1g.

      Page 175:  The paper's own summary of its conclusions.  “(a) That structurally and functionally the plica of man and Mammals differs entirely from the nictitating membrane of Birds.”  “(b) That there is reasonable ground for supposing that the plica of man and Mammals is morphologically different from the nictitating membrane.  But the morphology of the mammalian structures has yet to be worked out by a study of their ontogeny before any definite conclusion can properly be stated.” ↩ ↩ ↩

  2. 2.

    Wiedersheim, Robert.  The Structure of Man: An Index to His Past History.  Macmillan, London.  Translated by H. Bernard and M. Bernard.  Edited by G. B. Howes.  1895.  A translated textbook of comparative anatomy by Robert Wiedersheim, a professor at the University of Freiburg, that reads the structure of the human body as a record of descent from lower animals.  Its editor, G. B. Howes, suggested the translation for medical students and says the book is not an exhaustive treatise.  The article uses it for Wiedersheim's claims about the size and cartilage of the fold, and for the place of the fold on his list of vestigial organs.

    1. 2a.

      Page 148:  “Great interest attaches to the fold of the conjunctiva which lies at the median angle of the eye, and is known as the plica semilunaris (pl., Fig. 91).  This corresponds with the third eyelid, the so-called nictitating membrane, of the lower animals.  In Birds, Anurous Amphibians [some Sharks], and in many Reptiles it is highly developed, and, by means of a special muscular apparatus, can be drawn across the eyeball.  It serves not only to cover, but to keep clean the surface of the eye, the upper lid [which in Man performs that function] being immovable, and the lower slightly movable or but little developed.  In Man, as in the Apes, in association with the absence of a retractor bulbi muscle, this third eyelid has undergone great degeneration, but it may still enclose (more frequently in Negroes than in Caucasians) a cartilaginous support.” ↩ ↩ ↩

    2. 2b.

      Pages 148–149:  “Among sixteen pure Negroes this cartilage was found by Giacomini in twelve individuals.” ↩ ↩

    3. 2c.

      Page 149:  “The plica semilunaris varies greatly in size at different ages and in different races.  In the new-born child, and during the early years of life, it is broader than later, when it does not exceed 1½ to 2 mm. in breadth.” ↩ ↩ ↩

    4. 2d.

      Page 202:  In section B of the conspectus, under this heading:  “Retrogressively modified, the Organs having become wholly or in part functionless, some appearing in the Embryo alone, others present during Life constantly or inconstantly.  For the greater part Organs which may be rightly termed Vestigial”.  The entry for the fold reads:  “Plica semilunaris of the eye.” ↩

  3. 3.

    Gray, Henry.  Anatomy of the Human Body.  Lea & Febiger, Philadelphia.  20th American edition.  Edited by Warren H. Lewis.  1918.  A human anatomy textbook, in its 20th American edition, edited by Warren H. Lewis.  The article uses it for the standard description of the fold and of the caruncle beside it, and for an older report of smooth muscle in the fold.

    1. 3a.

      Page 1027:  “at the medial angle it forms a semilunar fold, the plica semilunaris.” ↩

    2. 3b.

      Page 1028:  “The caruncula lacrimalis is a small, reddish, conical-shaped body, situated at the medial palpebral commissure, and filling up the lacus lacrimalis.”  “Lateral to the caruncula is a slight semilunar fold of conjunctiva, the concavity of which is directed toward the cornea; it is called the plica semilunaris.  Müller found smooth muscular fibers in this fold; in some of the domesticated animals it contains a thin plate of cartilage.” ↩ ↩ ↩ ↩

    3. 3c.

      Page 1028:  The passage is about the caruncle.  “It consists of a small island of skin containing sebaceous and sudoriferous glands, and is the source of the whitish secretion which constantly collects in this region.” ↩

  4. 4.

    Arends, G., and U. Schramm.  “The structure of the human semilunar plica at different stages of its development: a morphological and morphometric study.”  Annals of Anatomy 186(3):195–207.  2004.  Two anatomists at the University of Lübeck who sectioned the fold in the eyes of eleven fetuses, two newborns and an old man, to see how it develops.  They found the fold packed with immune cells, blood vessels and mucus-making goblet cells, but with no cartilage, and with muscle cells only in a thin layer around some vessels.  The article uses them for the fold's immune role and for its lack of cartilage.

    1. 4a.

      Section Abstract:  “In this study development of the semilunar plica was examined histologycally by making sections through the eyes of eleven foetuses at different stages of gestation, two newborns and an old man.  We found that in the early stages of its development the semilunar fold covered a bigger part of the orbit and later did not keep up with the growth of the eyeball and the lids.”  "histologycally" is spelled that way in the original. ↩ ↩

    2. 4b.

      Section Abstract:  “Beside poorly differentiated buds of the surface epithelium which can be classified as rudiments of the nictitating or Harderian gland, serous glands were detected which could be beginnings of Krause's glands.” ↩ ↩

    3. 4c.

      Section Abstract:  “In general the palpebral side of the plica consisted of a higher number of epithelial layers and more goblet cells than its bulbar side.” ↩ ↩

    4. 4d.

      Section Abstract:  “In the tight collagenous connective tissue no elastic or reticular fibres or cartilaginous structures could be found.”  “Later on, some of them showed an enlarged lumen and were covered by a thin layer of muscle cells.”  “Muscle cells could not be detected.” ↩ ↩

    5. 4e.

      Section Abstract:  “A very dense concentration of leukocytes in the plica even in early intrauterine development was very striking and could be observed in this kind for the first time.”  “The discovery of dense infiltration with both specific and non-specific immune cells, abundant vascularization and secretory structures (goblet cells and surface enlargement) in the semilunar plica suggest that it plays an important role as a specialized organ in human eye protection.  Its anatomical position at the medial border of the eye supports this theory.  Its origin from the third eyelid of the mammals could be recognized to a certain extent.” ↩ ↩ ↩ ↩

    6. 4f.

      Section Abstract:  “Additionally, a new kind of plica gland was identified in which ductular structures, with an onion skin appearance, could be discriminated from mucous acini.” ↩ ↩

  5. 5.

    Walls, Gordon Lynn.  The Vertebrate Eye and Its Adaptive Radiation.  Cranbrook Institute of Science, Bloomfield Hills.  Hafner reprint, New York edition.  1942 (edition of 1963).  A book by Gordon Lynn Walls, a researcher in ophthalmology, written for general readers rather than specialists.  It explains how the eyes of backboned animals differ and why, by asking what each animal does with its eyes.  The article uses it for the structure of the third eyelid in birds and in mammals, and for the fold as a leftover of that eyelid.

    1. 5a.

      Page 38:  “In the inner canthus can be seen the plica semilunaris, a crescentic fold of conjunctiva which is a vestige of a third, sidewise-sweeping eyelid present in many animals, the nictitating membrane.  Neither the special muscles nor the special gland (Harder's) of the third eyelid are present, even as vestiges, in man.” ↩

    2. 5b.

      Page 420:  “A third lid, the vertical 'membrana nictitans', has consequently evolved as a fold of the conjunctiva at the inner or nasal corner of the lid opening (Fig. 142).  Being transparent, it can sweep the cornea from the nasal to the temporal side, to clean and moisten it, without shutting out the light.” ↩

    3. 5c.

      Page 420:  “In man, it is the heavy bony rim of the orbit, particularly the ridge bearing the eyebrow, which makes a retractor bulbi unnecessary.” ↩

    4. 5d.

      Page 424:  The passage is about birds.  “Its inner surface is always covered by an epithelium whose surface cells are built like unicellular feathers, which improve its cleansing action.” ↩

    5. 5e.

      Page 426:  “The nictitans has its ups and downs in the mammals.  Where it is well developed, it usually has a cartilaginous tarsus, but it never has a special musculature behind the globe as in lower vertebrates.  Hence, it slips over the eye only passively when the globe is slightly or markedly retracted.  Contrary to logical expectations, it is most rudimented in the lower mammals and has come back to greatest usefulness in some of the higher ones.  This probably explains the absence of its characteristic muscles, these having been discarded in early mammals to whom the nictitans was unimportant.  It is present in the duck-bill but lacking in the echidnas, and it is vestigial in rodents and others of the lower orders such as the insectivores, primates, and 'edentates'.” ↩ ↩ ↩ ↩ ↩ ↩

    6. 5f.

      Page 426:  “All carnivores except the skunk, whose eye protrudes greatly like that of a mouse, have a nictitans.  All can move it, though not all ever do so.  In only a few could it possibly be drawn all the way over the cornea, the 'haw' of the domestic cat being a familiar example of this rare degree of development.” ↩

    7. 5g.

      Page 427:  “Ungulates all have the nictitans, though with great interspecific variations, and usually with no apparent usefulness.  In the horse family, however, the nictitans is as extensive and as rapid in action as in many sauropsidans.  Its retention here is attributed to the need for special protection of the eye when feeding in deep grass”. ↩ ↩

  6. 6.

    Arao, Tatsuyoshi, and Edwin M. Perkins.  “The nictitating membranes of primates.”  The Anatomical Record 162(1):53–69.  Sep 1968.  A survey of the nictitating membrane, the third eyelid, in a sample of 22 non-human primate species, published in an anatomy journal in 1968.  In 21 of the 22 species the membrane was well developed, and only the chimpanzee's was vestigial.  The article uses it to test the claim that the membrane is vestigial across the primates, and to show that the human fold is not unusual among its nearest relatives.

    1. 6a.

      Section Abstract:  “Of 22 species of subhuman primates examined, 21 have a well developed nictitating membrane; it is vestigial only in Pan satyrus/troglodytes.  The sizes of membranes range from those that cover only one‐tenth of the eye (Gorilla gorilla) to those that extend over the entire eye (Perodicticus potto and Arctocebus calabarensis).  Nictitans, best developed in the lower primates, have a free border that is generally heavily pigmented.  With the exception of Galago crassicaudatus, all prosimians studied have a membrane that contains a cartilaginous band; cartilage is also present in Cacajao rubicundus, Cebus albifrons, and Saimiri sciureus.” ↩ ↩ ↩ ↩

    2. 6b.

      Section Abstract:  “The outer surface of the nictitans of all species of anthropoids has mucous glands; that of the prosimians has few of them.”  “Muscle fibers are present only in the nictitating membranes of Lemur monoz and Callithrix aurita.” ↩ ↩ ↩

  7. 7.

    Darwin, Charles.  The Descent of Man, and Selection in Relation to Sex.  Volume 1.  John Murray, London.  1871.  Charles Darwin's 1871 book on the descent of humans, here the first edition of its first volume.  The article uses it for Darwin's own words that the fold is "a mere rudiment" of the third eyelid, a claim about shared origin and reduction rather than about use.

    1. 7a.

      Page 23:  “The nictitating membrane, or third eyelid, with its accessory muscles and other structures, is especially well developed in birds, and is of much functional importance to them, as it can be rapidly drawn across the whole eye-ball.  It is found in some reptiles and amphibians, and in certain fishes, as in sharks.  It is fairly well developed in the two lower divisions of the mammalian series, namely, in the monotremata and marsupials, and in some few of the higher mammals, as in the walrus.  But in man, the quadrumana, and most other mammals, it exists, as is admitted by all anatomists, as a mere rudiment, called the semilunar fold.” ↩ ↩

  8. 8.

    Scadding, S. R.  “Do 'vestigial organs' provide evidence for evolution?”  Evolutionary Theory 5:173–176.  May 1981.  A critical paper by a zoologist at the University of Guelph on the argument that vestigial organs, which have no function, are evidence for evolution.  It traces the argument back to Darwin, Haeckel and Wiedersheim, whose list of eighty-six vestigial organs in the human body it says mostly have some function, and it concludes that vestigial organs give no special evidence for evolution.  The article uses it as the 1981 reply to the claim about the fold's function, which Naylor later answered.

    1. 8a.

      Page 175:  “The semilunar fold of the eye is simply that portion of the conjunctiva at the medial corner of the eye and as such aids in the cleansing and lubrication of the eye ball (Gray and Goss 1972, p. 1065).  The absence of a nictitating membrane in humans is of no significance to a discussion of the function of the semilunar fold.” ↩ ↩

    2. 8b.

      Page 176:  “Since it is not possible to unambiguously identify useless structures, and since the structure of the argument used is not scientifically valid, I conclude that 'vestigial organs' provide no special evidence for the theory of evolution.” ↩

  9. 9.

    Naylor, Bruce G.  “Vestigial organs are evidence of evolution.”  Evolutionary Theory 6:91–96.  Sep 1982.  A reply to Scadding's 1981 paper by a researcher in vertebrate paleontology at the University of Alberta.  He argues that a vestige need not be useless, since many vestigial organs keep a reduced or changed function, and that the human coccyx is one such organ.  The article uses it as the 1982 reply that counts the fold as evidence for descent.

    1. 9a.

      Page 92:  “Regardless of how definitions are altered to suit particular arguments, the fact remains that the coccyx, ear muscles, and semilunar fold of the eye provide powerful suggestive evidence for the evolution of man from "lower" primates.” ↩ ↩

  10. 10.

    Franks, Robert H.  “Vestigial organs: an idea destructive to medical science and our health.”  Creation 10(2):22–24.  Creation Ministries International.  Mar 1988 (edition of 2014).  A 1988 magazine article arguing that the idea of useless organs is destructive to medical science and to health, and that the plica has essential functions.  It was republished on the ministry's website on 3 December 2014.  The article uses it as an early creationist reply to the useless-organ claim, one that is right on the main point.

    1. 10a.

      Section Semi-lunar membrane:  “The plica semilunaris is a little fold of tissue at the inner corner of the eye.  Some evolutionists feel that it is a remnant of the third eyelid of a lower form, such as the third eyelid in birds and reptiles.  But in man this tissue has several essential functions.  If you did not have the tissue for support at that location, the eyeball would sink.  This would cause double vision.  The tissue not only supports the eyeball, but the tearduct as well.  Without this tissue, tears would drain over the cheeks causing a cosmetic problem.  This area also serves to collect foreign matter.” ↩ ↩

    2. 10b.

      Section Semi-lunar membrane:  “It is not true that this fold has no purpose.  It was designed.  And it does not represent the cleverly designed third eyelid of the bird which prevents the bird's eyes from drying out during flight.” ↩

  11. 11.

    Sanders, Lita.  “Badly designed arguments: 'vestigial organs' revisited.”  Creation Ministries International.  Nov 25, 2011.  A reply on a creationist ministry's website to a reader's request to answer another writer's list of supposedly useless human organs, taken one at a time.  The article uses it for the claim that the plica has a critical function for the health of the eye.

    1. 11a.

      Section 'Third eyelid' (plica semilunaris):  “The plica semilunaris actually has a critical function for the health of the eye.  It secretes the sticky substance that hardens into the crust we sometimes find around our eyes when we wake up.  This substance surrounds any particles that find their way into the eye, making them less likely to scratch the eye, and making it easier for the tears and eyelids to remove the particles from the eye.” ↩ ↩

  12. 12.

    Guliuzza, Randy J.  “Evolutionary Vestigial Features: Worse Than Myth, a Scam.”  Acts & Facts 54(5):16–17.  Institute for Creation Research.  2025.  An article in Acts & Facts, published by the Institute for Creation Research, that calls the vestigial-organs argument a scam and sets out the steps it says biologists follow.  The article uses it for that list, which names the plica among the obscure parts.

    1. 12a.

      “We can now begin to recognize the key steps of evolutionary biologists' vestigial organs scam.  Identify an obscure body part or action that most people have essentially no knowledge of (e.g., appendix, tailbone, adenoids, thymus and pineal glands, arrector pili muscles causing goosebumps, plica semilunaris or "third eyelid," etc.).”  “Declare that the part is a vestigial remnant of past evolution—since no one knows of a useful function.” ↩ ↩

  13. 13.

    Mitchell, Elizabeth.  “Vestigial Organ Arguments: Useful or Unnecessary?”  News to Know.  Answers in Genesis.  Oct 22, 2011.  A news commentary from Answers in Genesis that replies to a Fox News list of nine supposedly useless body parts, going through several of them.  It accepts that the plica is thought to be homologous with a third eyelid in some animals, but says it is far from useless.  The article uses it because Answers in Genesis accepts the homology and disputes only the claim of uselessness.

    1. 13a.

      “The #1 item—the plica semilunaris (third eyelid)—according to Fox News, is "left over from what's known as a 'nictitating membrane,' which is still present in animals like chickens, lizards, and sharks."  While this fold of protective eye covering is thought to be homologous with the more complete protective covering in some animals, it is far from useless.  Secretions from this "#1 useless body part" mop debris from your eyes.” ↩ ↩

    2. 13b.

      “Similarity of parts— homologies— between different creatures demonstrates that our common Designer used well-engineered designs to serve various functions.” ↩

  14. 14.

    Knop, Nadja, and Erich Knop.  “Conjunctiva-associated lymphoid tissue in the human eye.”  Investigative Ophthalmology and Visual Science 41(6):1270–1279.  2000.  A histology study of 53 complete conjunctival sacs and eight lacrimal sacs from donor eyes, prepared as whole mounts and sections.  It found diffuse lymphoid tissue in every specimen and organized follicles in about three fifths, mainly in the lid lining, and named the system conjunctiva-associated lymphoid tissue.  It shows that the immune cells in the plica are part of the eye surface's working immune system, and also that the fold is not its main site.

    1. 14a.

      Section Abstract, Results and Conclusions:  “In all specimens, diffuse lymphoid tissue of lymphocytes and plasma cells, most of which were IgA positive, formed a thin layer in the lamina propria.”  “In approximately three fifths of the conjunctival sacs, organized follicular accumulations were embedded in this layer.”  “Human conjunctiva and lacrimal drainage system show an associated lymphoid tissue (suggesting the term conjunctiva-associated lymphoid tissue [CALT]) that contains all components necessary for a complete immune response.” ↩

    2. 14b.

      Section Abstract, Results:  “Lymphoid tissue was mainly observed in the palpebral conjunctiva, more pronounced in the upper than in the lower lid.”  The palpebral conjunctiva is the part lining the inside of the eyelids. ↩

  15. 15.

    Sisson, Septimus.  The Anatomy of the Domestic Animals.  W. B. Saunders, Philadelphia.  2nd edition.  1914.  A book on the anatomy of domestic animals.  The article uses its description of the horse's third eyelid to show how the mechanism works.

    1. 15a.

      Page 859:  The description is of the horse.  “The third eyelid (Palpebra tertia) is situated at the medial angle of the eye.  It consists of a semilunar fold of the conjunctiva, known as the membrana nictitans, which covers and partly encloses a curved plate of hyaline cartilage.”  “Numerous minute lymph nodules occur in the membrana nictitans, and the deep part of the cartilage is surrounded by a gland which resembles the lacrimal gland in structure”.  “Ordinarily the third eyelid extends very little over the medial end of the cornea, but when the eyeball is strongly retracted, the membrana is protruded over it so as to measure about an inch (ca. 2-3 cm.) in its middle.  This effect results from the pressure of the eyeball and its muscles on the fat which surrounds the deep part of the cartilage.” ↩ ↩ ↩

  16. 16.

    Thompson, J. W.  “The nerve supply to the nictitating membrane of the cat.”  Journal of Anatomy 95(Pt 3):371–385.  Jul 1961.  A laboratory study of the nerves that reach the cat's nictitating membrane, its third eyelid, from the sympathetic nervous system.  Thompson traced them in twelve dissected preparations and confirmed them by stimulating the nerves in five cats prepared as spinal animals.  He found that the medial and inferior smooth muscles each have their own sympathetic nerve, which reaches it along branches of the trigeminal nerve.  Thompson worked at the Institute of Basic Medical Sciences, Royal College of Surgeons, London.  The article uses it for the two muscles that pull the cat's membrane back, the counterpart it asks about in the human fold.

    1. 16a.

      Page 371:  “The nictitating membrane of the cat is commonly used to study sympathetic transmission”.  “The membrane contains two thin sheets of smooth muscle.  Acheson (1938) showed that these smooth muscles, which he named the medial and inferior muscles, arise deeply in the orbit from the fascial coverings of the medial and inferior recti muscles, respectively, to be inserted into adjacent sides of a T-shaped piece of cartilage (Text-fig. 1 a, b).”  A figure interrupts the sentence after "medial and inferior muscles,". ↩

    2. 16b.

      Page 384:  Reference list.  “PATON, W. D. M. & THOMPSON, J. W. (1953).  The muscles retracting the cat's nictitating membrane.  J. Physiol. 120, 55P.” ↩

  17. 17.

    Rehorek, Susan J., and Timothy D. Smith.  “The primate Harderian gland: does it really exist?”  Annals of Anatomy 188(4):319–327.  2006.  A microscope study of the orbits of newborn primates from twelve species, lemur and loris groups among them, and tarsiers, marmosets and their relatives.  The Harderian gland, a gland in the front of the eye socket, was present in numerous species at birth, and its size ranged from large to absent, with tarsiers the group that lacks it.  The authors say its role needs further study.  The article uses it to correct the claim that the gland is absent in humans, since it is present in the human fetus and newborn.

    1. 17a.

      Section Abstract:  “The Harderian gland, an anterior orbital structure, is either absent or vestigial in primates.  This is based upon gross anatomical observations of scattered adult specimens.  Though largely absent in the adult human, it is present in the fetal and neonatal stages.” ↩ ↩

    2. 17b.

      Section Abstract:  “The relative size of the neonatal primate Harderian gland can be subdivided into five separate categories, ranging from large to absent (tarsiers), with no apparent phylogenetic trends.” ↩ ↩

  18. 18.

    Coyne, Jerry A.  Why Evolution Is True.  Viking, New York.  2009.  A book about evolution by Jerry A. Coyne.  The article cites it only for his definition of a vestige: a trait can be vestigial and functional at the same time.

    1. 18a.

      Page 58:  “Evolutionary theory doesn't say that vestigial characteristics have no function.  A trait can be vestigial and functional at the same time.  It is vestigial not because it's functionless, but because it no longer performs the function for which it evolved.” ↩ ↩

  19. 19.

    Bergman, Jerry.  “Do any vestigial structures exist in humans?”  CEN Technical Journal 14(2):95–98.  2000.  An article arguing that the list of vestigial organs in humans has shrunk from 180 in 1890 to 0 in 1999.  It rejects the newer "revisionistic" definition of a vestigial structure as too vague, since it would label almost every structure as vestigial.  The article uses it as the source that lists the plica among the 180 structures once called vestigial.

    1. 19a.

      Page 96:  The caption to a figure listing "plica semilunaris" among other structures.  “A list of some of the 180 structures that were considered vestigial in the early 1900's.  It is now almost unanimously agreed that most of these structures have at least one function.” ↩

  20. 20.

    Sarfati, Jonathan, and Don Batten.  “'Vestigial' Organs: What do they prove?”  Creation Ministries International.  Feb 14, 2006.  A web article by two authors, published by Creation Ministries International, on what vestigial organs prove.  The article uses it as the reply from this literature that could meet the modern definition of a vestige.

    1. 20a.

      “Second, even if the alleged vestigial organ were no longer needed, it would prove devolution not evolution.  The creation model allows for deterioration of a perfect creation.” ↩

  21. 21.

    Heralgi, Mallikarjun et al.  “Persistent unilateral nictitating membrane in a 9-year-old girl: a rare case report.”  Indian Journal of Ophthalmology 65(3):253–255.  Mar 2017.  A case report of one nine-year-old girl with a membrane covering one-third of the cornea of one eye, which was removed by surgery.  The authors say it is only the second human case of a persistent nictitating membrane that they found in the literature.  The article uses it as the modern human case of a full third eyelid, and says it adds little either way.

    1. 21a.

      Section Case report:  “a translucent membrane arising from medial canthus and spreading out in a horizontal fashion over the bulbar conjunctiva and covering one-third of the cornea in the primary gaze”.  “The dimensions of the membrane were 20 mm horizontally and 18 mm vertically”. ↩

    2. 21b.

      Section Case report:  “Histopathological report showed stratified squamous epithelium with underlying loose areolar connective tissue and nonspecific inflammatory cells around blood vessels.  No elastic, reticular fibers, cartilaginous structures, or muscle cells could be found”. ↩ ↩

    3. 21c.

      Section Discussion:  “The presence of nictitating membrane in humans is rare and only one such case report by García-Castro and Reyes de Torres was found in the literature where they describe a newborn with features of trisomy 18 syndrome having bilateral nictitating membrane.” ↩