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Adv Physiol Educ 42: 90–98, 2018;

doi:10.1152/advan.00161.2017.

A PERSONAL VIEW

Using Hawkeye from the Avengers to communicate on the eye
X Barry W. Fitzgerald
Intensified Reaction & Separation Systems, Department of Process & Energy, Delft University of Technology, Delft,
The Netherlands
Submitted 26 October 2017; accepted in final form 6 December 2017

Fitzgerald BW. Using Hawkeye from the Avengers to communi- whereas Hawkeye, or Clint Barton, who also possesses many
cate on the eye. Adv Physiol Educ 42: 90 –98, 2018; doi:10.1152/ of Ramonff’s abilities, is an expert archer.
advan.00161.2017.—Superheroes, such as Iron Man, Captain Amer- In my opinion, Hawkeye is one of the more grounded
ica, Wonder Woman, Batman, and Hawkeye, have appeared in nu- superheroes. He has not been the subject of an experimental
merous films, displaying their range of incredible superpowers and
abilities. Therefore, it is unsurprising that many people would not only
treatment and does not wear an Iron Man-like suit. Just like the
wish to attain these powers, but also to learn about scientific acces- average person, Hawkeye fatigues and must eat to replenish
sibility to these powers. Popular culture characters such as superhe- his energy reserves. For this alone, he is a superhero to
roes can provide a unique platform for the communication of difficult whom the student can relate. Hawkeye’s abilities as an
scientific concepts. In the classroom, these characters can be used to archer can be attributed to his exceptional eyesight, which
communicate learning objectives to students in an interesting, fun, and can act as an innovative platform to introduce the physiol-
accessible manner by taking advantage of student familiarity with the ogy and anatomy of the eye to students (13). The character
characters. Hawkeye, a member of the Avengers, is one such super- can also be used to motivate discussion on how current
hero who can be utilized by educators. His powers can be attributed scientific research could be used in the near future to
in part to his advanced eyesight, which has physiological aspects in increase the acuity of human eyesight (13).
common with many birds of prey. Hence, Hawkeye can instigate
discussion on the physiology of the human eye, while also allowing
Using superheroes such as Hawkeye as a platform for the
for comparison with other species, such as birds of prey, and reflection communication of learning objectives related to physiology
on advancements related to genetic engineering and wearable tech- can have a multitude of benefits for educators and students (3,
nologies. In addition, in my experience, Hawkeye has proven to be a 61, 62). First, superheroes can be used to motivate the student’s
highly suitable popular culture character for use in scientific commu- initial interest in a new topic or subject. Given the popularity of
nication and outreach. superhero films, it is likely that the majority of students will
education; eyesight; popular culture; scientific outreach; superheroes
have seen, or, at the very least, be aware of the superhero
character. Second, the use of superheroes in the classroom
represents a departure from traditional analogies used to en-
force or support the delivery of learning objectives. Third, the
INTRODUCTION topic of superheroes can be used to inspire educators in the
design of laboratory experiments, in-class assignments, student
In recent years, there has been an upsurge in the popularity projects, and final examination questions. The use of superhero
of the superhero genre, motivated in part by the large number paradigms can also foster stronger links between educators and
of successful superhero films. Some films focus on superhero students, and between physiological aspects of the human body
teams, such as the X-Men and the Avengers, with team and the world of the student. In addition, Hawkeye can be used
members having a diverse range of complementary abilities. In in scientific outreach to help bridge the gap between academic
the Avengers, Iron Man (Tony Stark) wears a powered exo- research and the general public.
skeleton suit that provides him with increased strength and the In this article I will discuss the physiology of Hawkeye’s
ability to fly, whereas Captain America (Steve Rogers) is a eyes, drawing comparisons with the vision of birds of prey. I
World War II super-soldier whose physiology has been per- will motivate Hawkeye using one scene from the 2012 film The
manently altered following treatment with an experimental Avengers (2). In addition, I also briefly suggest how Hawkeye
serum and radiation exposure (3). Other team members are can be used to motivate other topics, such as genetic engineer-
aliens, such as Thor, and the Vision is an android powered by ing and transgenesis. Finally, I propose pedagogical applica-
an alien relic known as an Infinity Stone. Nonetheless, the tions of Hawkeye in the classroom and suggest methods for
abilities of some Avengers are firmly based on genetics and motivating classroom discussions as well as planning experi-
human physiology. Black Widow, or Natasha Romanoff, is a ments connected to learning objectives.
highly trained Russian spy with Olympic athlete level fitness,
agility, and reflexes and an expert in hand-to-hand combat,
Hawkeye: the Superhero
Many people will be familiar with Hawkeye, as portrayed by
Address for reprint requests and other correspondence: B. W. Fitzgerald, Jeremy Renner, from films such as The Avengers (2), Aveng-
Delft University of Technology, Process & Energy Department, Intensified
Reaction & Separation Systems, Building 34K, Room 34K-0-100, Leeghwa- ers: Age of Ultron (1), and Captain America: Civil War (4).
terstraat 39, 2628 CB Delft, The Netherlands (e-mail: barry.w.fitzgerald The character first appeared in the Marvel comic book Tales of
@gmail.com; b.fitzgerald@tudelft.nl). Suspense, no. 57, in 1964 (29), and he joined the Avengers in
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HAWKEYE EYE 91

Avengers, vol. 1, no. 16, from 1965 (28). His two key physi-
ological characteristics are his speed of reaction and superhu-
man eyesight, which both contribute to his ability as a master
archer. To rapidly fire one accurate arrow after another, Hawk-
eye must be at the peak of human conditioning, with highly
developed tendons and ligaments, as well as exemplary eye-
hand coordination. However, the focus of this article is solely
on his vision.
To motivate this topic, I will refer to one particular scene
from the 2012 film, The Avengers (2). In the film, the Avengers
battle the Chitauri, an alien army, that arrives via a wormhole
above Stark Tower in New York City. During the fight scene,
Hawkeye is perched on a skyscraper firing arrows at Chitauri
gliders flying through the New York City skyline. At one point,
Black Widow flies an alien glider with Loki, the main antag-
onist of the film, in pursuit. Sensing the danger, Black Widow
Fig. 2. The human eye with the cornea, lens, optic nerve, retina, and fovea all
contacts Hawkeye for assistance. Hawkeye calmly loads an indicated.
arrow in his bow and tracks the gliders of Black Widow and
Loki. The camera then zooms in on the pair of gliders, and, in
doing so, gives the audience an impression of Hawkeye’s ture, and increased photoreceptor density in the eyes. I will also
advanced vision. Hawkeye confidently states “I got him” and compare the position and associated visual field of his eyes
fires an arrow at Loki. Although Loki catches the arrow, it with those of birds of prey.
explodes, destroying Loki’s glider and allowing Black Widow Accommodative mechanism. First, I will consider the accom-
to escape. modative mechanism in the human eye (Fig. 2). The human
eye is sensitive to electromagnetic waves, with wavelengths
Hawkeye and Birds of Prey ranging from 390 nm up to 700 nm, extending from violet to
red. When light enters the eye, it is refracted by the cornea and
Hawkeye’s accuracy with the bow and arrow is dependent the lens to focus the light onto the retina, the light-sensitive
on his eyesight, which is more advanced than the average part of the eye, where the light excites photoreceptor cells,
human eyesight. As indicated previously, Hawkeye has not leading to chemical and electrical nerve impulses (Fig. 2).
been enhanced via experimentation, technology, or alien arti- Neurons, known as retinal ganglion cells, collect the impulses
facts. He has not been administered an experimental serum like and transmit them via the optic nerve to the brain. The cornea
Steve Rogers (3), and he has not been genetically modified has a fixed spherical shape, acts as a converging lens, and
after exposure to cosmic rays like the Fantastic Four. Hence, I accounts for roughly two-thirds of the refractive power of the
must conclude that he has inherited the genetic propensity for eye. The ciliary muscles surrounding the lens change the
advanced eyesight from his parents. The nickname “Hawkeye” refractive power of the lens, a process known as accommoda-
reflects this and suggests that he has many attributes in com- tion. To resolve distant objects, the lens will assume a thin and
mon with birds of prey (Fig. 1). There are several aspects of the flat geometry, whereas, for closer objects, the lens is thicker
eye that could be central to Hawkeye’s vision. For brevity, I and rounder.
will highlight what I believe are three key aspects, which are The accommodative process in birds of prey is different than
advanced accommodative mechanisms, differing retinal struc- for humans (23, 60) (Fig. 3). While the ciliary muscles con-
nected to the lens change the refractive ability of the transpar-
ent lens in a human eye, accommodation is facilitated in birds
of prey through the deformation of both the lens and cornea.
Corneal accommodation plays a key role in the overall vision
of birds. In front of the cornea is a layer of air, and behind the
cornea is the aqueous humor, which is a waterlike liquid. With
increased curvature, the cornea has greater refractive power
and can thus exploit to a greater extent the interface between
the air and aqueous humor, which have differing refractive
indexes. This process is facilitated by the anterior sclerocorneal
muscles, which are striated or striped muscles in the eye.
The refractive power of an optical device is a measure of the
degree by which it bends light and is calculated from the
reciprocal of the focal length (P ⫽ 1/f). It is measured in units
of diopters (D) or m⫺1. For a human eye, the refractive power
is ~60 D, with ~40 D due to the refractive power of the cornea.
While the refractive power of the eyes in birds of prey is only
slightly larger, the corneal and lenticular accommodative pro-
Fig. 1. Sketch of the eye of a bird of prey. [Image credit: Jean-Philippe Frimat. cesses allow for greater flexibility and adaptivity for focusing
From Fitzgerald (13) with permission.] images on the retina. In addition, some avian species can be

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92 HAWKEYE EYE

Fig. 3. An example of the anatomy of an avian
eye. [From Whittow (60) with permission from
Elsevier.]

emmetropic and myopic at the same time (23). This means that capture prey in the momentum phase of a dive or when the
a bird can simultaneously focus on two objects: one that is gannet first enters the water. Second, prey can be acquired
distant and one that is closer to the bird. Such accommodative during the wing-flapping phase, where the gannet pursues the
mechanisms can explain why Hawkeye is able to track multiple prey through the water. An experimental study of plunging
targets at varying distances in the film The Avengers (2). gannets suggests that they switch from aerial vision to clear
While Hawkeye temporarily retires from the Avengers in underwater vision in ⬍0.1 s (8, 32). The study suggests that the
Avengers: Age of Ultron (1) and returns to action in Captain clear aquatic vision is due to the reshaping of the lens against
America: Civil War (4), he will eventually have to permanently the iris, which controls the amount of light entering the eye.
retire due to degradation of his accommodative abilities. In a When underwater, the ciliary muscles pull the lens forward
normal human eye, the process of accommodation deteriorates rather than deforming the lens in the traditional manner. The lens
over time, leading to the condition of presbyopia, a decrease in then slightly protrudes beyond the iris and increases the refractive
the eye’s ability to focus on close objects. While this condition ability of the lens. For Hawkeye, this process may be sufficient
will not affect Hawkeye’s ability to focus on distant objects to correct for the loss of refractive power of the cornea while
and could be corrected with advanced surgical techniques (14), underwater and allow him to retain his advanced accommoda-
Hawkeye will certainly suffer some loss of corneal and lentic- tive mechanism.
ular accommodation with age, just like many avian species Photoreceptor cells and foveae. While Hawkeye’s accom-
(23). In fact, Hawkeye is shown as blind in the Old Man Logan modative mechanisms are crucial for focusing light onto his
comic book (37), where it appears that he has developed retina, his visual acuity is also dependent on his retinal struc-
cataracts in his eyes and suffered serious eye trauma rather than ture. In the human eye, there are two main types of photore-
experienced degradation of his accommodative mechanisms. ceptor cells, referred to as rods and cones. Four genes are
We have yet to see Hawkeye using his trusted bow and associated with these cells: one gene for rods and three genes
arrow underwater in the Marvel Cinematic Universe films. The for cones. The RHO gene leads to the production of the protein
human eye loses the refractive ability of the cornea when rhodospin, which is found in the rods. The other three genes are
underwater, because the refractive index of water is approxi- OPN1SW, OPN1MW, and OPN1LW, which encode for the
mately the same as the refractive index of the fluids in the eye. blue-sensitive opsin protein, green-sensitive opsin protein, and
As a result, the lens is unable to compensate for the missing red-sensitive opsin protein, respectively (40). Here SW stands
refractive power, leading to the projection of an image beyond for short wave, MW is medium wave, and LW is long wave.
the retina, known as hyperopia. This explains why someone The breakdown of cones in the eye is ~65% red cones, 33%
sees blurry images when underwater. If Hawkeye were trying green cones, and 2% blue cones, where the blue cones have the
to incapacitate a villain underwater, he would need additional greatest sensitivity (48). Hence, humans, like cats (50), have
accommodative mechanisms from certain birds of prey to trichromatic vision, given that we possess cones sensitive to
retain visual acuity. The gannet is a bird of prey that seemingly three wavelength ranges. Rods provide lower resolution, but
retains visual acuity when moving from air to water. Gannets they are very sensitive to lighting conditions. Thus they are
are large seabirds that search for fish from above the water important for vision in poorly illuminated conditions. At the
before plunging into the water after the prey. They can capture lowest luminance, where the rods are primarily active, vision is
prey during two different phases of the dive. First, they can referred to as scotopic. On the other hand, cones allow for high

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HAWKEYE EYE 93

spatial resolution or visual acuity, while being insensitive to
light intensity. For indoor lighting or sunlight, cones mainly
contribute to variances in vision, since the contribution from
the rods saturates, a vision referred to as photopic (48).
In the average human eye, rods and cones are located in
differing parts of the retina (22, 48). The human eye comprises
between 90 and 120 million rods and, on average, 4.5 million
cones. Cones are packed into the fovea centralis in the center
of the macula lutea in the retina, whereas rods are found in
other regions of the retina. In an investigation on the photore-
ceptor count in eyes from 21 donors, the density of cones in the
fovea region has been shown to be in excess of 125,000
cones/mm2, whereas the density of rods peaks somewhere in a
ringlike area 3–5 mm from the fovea at 150,000 rods/mm2
(22). In comparison, an eagle can have up to 1,000,000 cones/
mm2 in its fovea (10).
In recent years, there has been a notable advancements in
relation to noninvasive medical imaging of microvascular flow
in the vicinity of the skin (25, 34, 35, 44), imaging of bowels
(31), and the imaging of the human eye (21, 46, 59), as well as
the eyes of birds of prey (51). For the latter two cases, the
Fig. 4. Relative receptor density for an ideal bird of prey. [From Tucker (55)
noninvasive approach means that the retinal structure of both with permission.]
birds of prey (51) and humans (59) can be studied in a manner
that is both accurate and safe for the subject. One approach that
is ideally suited for ophthalmic imaging, such as retinal cross- As he carefully tracks one glider, he appears to focus on a
sectional imaging, is optical coherence tomography (OCT) (21, second glider at the same time. Without checking on the
46, 51). Similar to ultrasonic pulse-echo imaging, OCT is a position of the first glider, he fires the arrow, hits the target and
technique that has been developed to facilitate noninvasive then prepares to fire at the second glider. As mentioned in the
cross-sectional imaging in biological systems. OCT uses inter- previous section, this multitargeting capability may also be
ferometry to generate a two-dimensional image of tissue using facilitated by the combination of corneal and lenticular accom-
back-reflected photons from tissue of low-coherence light, with modative processes. Nonetheless, this multitracking example
a wavelength of 830 nm. Three-dimensional images of the offers support for the proposal that Hawkeye is bifoveate.
retinal structure for two diurnal hawks using OCT show that The positions of Hawkeye’s foveae would be markedly
diurnal hawks are bifoveate, meaning that they possess two different from birds of prey as influenced by the skull structure.
foveae in the retina, classified as the deep fovea and shallow For birds of prey, the deep fovea has a line of sight that is
fovea (51). Just like the human eye, these foveae contain cones greater than 30° to the side of the reference line through the
and are associated with acute vision. The deep fovea is located skull (Fig. 5). On the other hand, the shallow fovea makes an
in the nasal or central region, which is close to the nasal part of angle of ~15° with the same reference line. These angles are
the bird’s skull, whereas the shallow fovea is found in the dependent on the species and on the extent to which the eye can
temporal region, which is closer to the temple of the bird (see rotate in the socket, as many avian species have limited
Fig. 3). Therefore, the retinal structure in birds of prey is movement of the eye in the socket (23). As a result, a bird of
markedly different from the retinal structure of the human eye. prey needs to move its head in order for the light from an object
Of the two foveae, the deep fovea is more clearly discernible of interest to fall on the deep or shallow fovea. For objects at
and developed. For a Broad-winged Hawk, the depth of the distances ⬎20 m, one study has shown that birds of prey spend
deep fovea has been shown to be 66% of the total retinal most of their time looking at the objects sideways in order for
thickness (51), whereas the shallow fovea is imaged as a slight the light to fall on the deep fovea, the region with the highest
depression in the retina. Conversely, retinal images of another visual acuity (55), while a recent study has also shown that
species, the Short-tailed Hawk, clearly demonstrate the pres- birds of prey, such as the Northern Goshawk and Red-tailed
ence of the shallow fovea. This suggests that the extent of the Hawk, search for prey using random head turns (42). Given
shallow fovea could depend on the species or on an individual that his eyes can move in their sockets, it is unlikely that
from that species (51). Typically, the central fovea contains a Hawkeye has to significantly move his head when focusing on
higher density of photoreceptor cells in comparison to the targets using either his supposed deep or shallow fovea. Birds
shallow fovea (Fig. 4). of prey do not have the same luxury. When facing forward,
As shown in the OCT study of Ruggeri et al. (51), some light from an object of interest will predominantly fall on the
individuals in avian species do not have a noticeable shallow shallow fovea (Fig. 5). While this is in the binocular vision
fovea. It may very well be the case that Hawkeye does not have field, it does not provide the bird with the greatest visual acuity.
a shallow fovea, given that the human eye ordinarily does not Unlike birds of prey, Hawkeye does not appear to search for
have such a fovea. However, there is credible evidence from prey in a random fashion. In The Avengers (2), Hawkeye scans
the film The Avengers (2) that suggests the contrary. At one his surroundings in a seemingly methodical fashion, which
instance in the Battle of New York City, we find Hawkeye may indicate that his bifoveate eyes make similar angles with
perched on a skyscraper with an arrow loaded and ready to fire. the reference line of his skull. Given the physiology of his eyes,

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94 HAWKEYE EYE

interacting with the photosensitive component of the cone. The
droplet, of which there are six different types, acts as a filter for
different wavelengths, meaning that the response of each cone
is dependent on the cone type and the type of oil droplet. Thus
birds of prey can process a much larger range of colors in
comparison to the human eye (23). From the films, thus far,
there is no evidence to suggest that Hawkeye can detect UV
wavelengths through the presence of a fourth cone. To
strengthen the association with birds of prey, it might be an
idea for writers of future Marvel Cinematic Universe films to
reveal that Hawkeye can “see” or detect UV light.
Size, position, and visual fields. I now reflect on the size,
position, and visual field of the human eye in comparison to
birds of prey. One of the most striking aspects of the anatomy
of birds of prey is the size of their eyes in comparison to their
skull. In the case of humans, the eyes take up ~5% of the
volume of the skull, whereas for many birds of prey, the eyes
occupy ~50% of the volume of the skull (23). The position of
the eyes in humans and birds of prey is also noticeably
different. Humans have front-facing eyes, whereas the eyes of
many birds of prey are angled away from the midline or
reference line of their skulls (Fig. 5).
Fig. 5. An illustration of the frontal section of the head of an idealized bird of Birds of prey and humans have markedly different visual
prey, such as a raptor, showing the positions of the deep and shallow foveae. fields. Figure 6 shows a comparison of the horizontal visual
The line of sight (LOS) through the pupil to the deep fovea is shown in the fields of a Red-tailed Hawk and a human. For the Red-tailed
diagram. [From Tucker (55) with permission.]
Hawk, this is the visual field when their eyes are fully con-
verged or fully forward toward the reference line of the skull
eye rotation in his sockets is apparently not an issue. If he (Fig. 5). The Red-tailed Hawk has a binocular field of ~33°,
possesses both a deep and shallow fovea, unlike some birds of which extends 16.5° left and right of the skull reference line.
prey, Hawkeye can rely on both eye and head rotation to allow The lateral or monocular field, shown in white in the figure,
light for an object to fall on either the shallow or deep fovea. extends a further 122° on either side of the binocular field,
For Hawkeye to possess the visual acuity suggested in the “I while their blind spot makes up the remaining 82°, which is
got him” scene from The Avengers (2), he would certainly need more or less directly behind them. The visual field is species
a deep fovea similar to that of birds of prey, a photoreceptor dependent, with some having larger or smaller binocular fie-
density similar to that of birds of prey and advanced avian-like lds and blind spots. For example, the species Cooper’s Hawk
accommodative mechanisms outlined in the previous section. has a binocular field of 36° and a blind spot of 66° (45). For the
In the scene, Hawkeye is perched on a building looking down human eye, the binocular field is ~120° with a monocular field
East 42nd Street toward the East River. At the moment that his of roughly 35° either side of the binocular field. A blind spot of
vision is shown to zoom in on the gliders of Black Widow and 170° makes up the rest of the human visual field, which is over
Loki, the gliders are ~700 m from Hawkeye’s position. From twice the size of the blind spot for the Red-tailed Hawk.
the film, I estimate that the gliders are ~3 m in length and 1.5 Experiments have found that some birds of prey spend more
m in width. The golden eagle is reported to be able to instantly time looking sideways at objects when the distance to the
adjust its vision to zoom in on a rabbit with a length of up to object is larger than 20 m, which indicates that, to use the acute
0.5 m that is a distance of more than 3 km from the eagle (15).
Therefore, the advanced vision and automatic zooming ability
of his eyes attributed to Hawkeye in this scene from The
Avengers (2) is representative of the ability of birds of prey.
Finally, in this section, I will reflect on the structure and
additional cone in the eyes of birds of prey and potentially
possessed by Hawkeye. Many avian species, including hawks,
are tetrachromats, meaning that their retina contains four dif-
ferent cones, where the additional cone is sensitive to ultravi-
olet (UV) wavelength. UV vision has been shown to be
important for tracking the scent marks of prey while hunting
(19). For example, scent marks deposited by voles have been
shown to have a UV spectra (57). Another study has shown
that the UV cones are magnetoreceptors, used by certain
species to enable orientation with the Earth’s magnetic field Fig. 6. Comparison of the horizontal visual fields for a Red-tailed Hawk and
a human. The binocular field is shaded in gray, the lateral field is in white, and
(41). The structure of cones in birds of prey is further compli- the blind spot is in black. For the Red-tailed Hawk, this is the visual field when
cated by the presence of oil droplets in the cone. When light the eyes are converged toward the frontal part of the skull or closer to the
interacts with a cone, it passes through the oil droplet before reference line (see Fig. 5).

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HAWKEYE EYE 95

vision of the deep fovea, birds of prey look at objects using a efficients for nonspherical particles that experience drag and
single eye and the monocular field of vision (55). In a recent lift forces in industrial processes (33, 36).
study involving a species known as Harris’s Hawks, the birds
displayed a high frequency of head movements, and hence Creating Hawkeye’s Eyesight
sideways head positions, when selecting a target, which may be
associated with their increased dependence on their monocular From the outset of this article, I have argued that the only
vision field (47). feasible biological explanation for Hawkeye’s advanced eye-
Given his appearance, it is clear that Hawkeye, and the actor sight is thousands of years of genetic evolution in the form of
Jeremy Renner who plays the character for that matter, cannot adaptation, genetic drift, or mutation of his DNA. This sug-
have the same visual field of a bird of prey such as the gests that members of Barton’s family also have similar eye-
Red-tailed Hawk. Hawkeye’s eyes are front facing in his skull sight. In the Marvel comic books, this is the case, as Clint’s
and certainly not angled away from the midline of his skull. brother, Barney Barton, is also an accomplished archer, thanks
Even if Hawkeye is bifoveate, possesses a larger density of to his augmented vision. While evolution could lead to the
photoreceptor cells, and has advanced accommodative mech- emergence of Hawkeye-like sight in the human species in the
anisms, his only recourse to overcome the limitations presented future, although it is highly unlikely, we may have to wait
by the size of his blind spot is to rotate his head. In The millennia and many generations for these genetic changes to
Avengers (2), Hawkeye can be seen continually moving his take hold. The genetic mutation of the Barton DNA to allow
head as he assesses the attack formation and flight paths of the for the family’s advanced vision is associated with precise
Chitauri gliders in New York City, whether perched above the changes of the nucleotides, the subunits of DNA, in the
city on a skyscraper or engaging targets from a ground posi- double-helix structure of the DNA. The three primary means of
tion. Hawkeye’s head movement is a necessity as he tries to instigating mutation in DNA is by changing of a nucleotide,
mimic the visual field of a bird of prey. In one scene, it appears deletion of a nucleotide, or the insertion of a nucleotide
that Hawkeye takes down a glider behind him without looking between two existing nucleotides. Once the DNA changes, it
or turning his head. However that scene takes place after a can be passed onto the next generation via genetic inheritance
verbal exchange with Iron Man during which the focus of the (13). For example, Barton’s DNA, just like our DNA, contains
film moves from Hawkeye on the rooftop to Iron Man flying the genes OPN1SW, OPN1MW, and OPN1LW behind the
down a New York street. The film focuses on Iron Man for production of the blue, green, and red photoreceptor cones in
approximately 4 s, which would have been sufficient time for his foveae. For Barton to have the visual acuity of birds of
Hawkeye to quickly look behind himself, identify a target, and prey, he will certainly need a density of these cones compara-
then use his experience as an archer to predict the correct ble to birds of prey in his foveae. The production of photore-
moment to blindly fire the arrow at the glider. ceptor cones are regulated by transcription factors (TFs), which
Just like birds of prey, head movement is important for are proteins that control how often a gene is transcribed into a
Hawkeye to identify and focus on targets. Hawkeye normally protein. In the case of the human eye, the TFs for photoreceptor
uses his advanced eyesight to isolate a target from a stationary cones include OTX2, NR2E3, and CRX (54), whereas, in birds,
position and then fire an arrow at the target, such as in The the TFs will be slightly different (7). Thus the TFs for birds are
Avengers (2). Thus it is rare that Hawkeye will be traveling at in some way linked to the larger density of cones, in compar-
large speeds while firing arrows. Nevertheless, head position is ison to humans, in their foveae.
crucial for birds of prey when engaging a prey. As shown by Rather than wait for the TFs in human DNA to mutate and
Tucker (55), birds of prey tend to hold their head in a sideways change over many generations to match those of birds, we
position when viewing a prey that is straight ahead. Birds of could speed up the process by using genetic engineering
prey keep their head in this position to ensure that light from approaches. The field of genetic editing and DNA site-specific
the target falls on the deep fovea, thus providing maximum modification has been reinvigorated recently following the
visual acuity. However, in maintaining a sideways position, the development of the genetic engineering tool CRISPR/Cas9
bird experiences increased aerodynamic drag. Tucker suggests (11, 13, 20, 27, 49, 53). CRISPRs, which stands for clustered
that a bird can follow a logarithmic spiral flight path to regularly interspaced palindromic repeats, are DNA chains of
minimize the effects of aerodynamic drag and still keep its base sequences separated by gaps in a genetic sequence,
head sideways during flight (55). Despite having to fly a longer whereas Cas9 is a restriction endonuclease or enzyme that can
path, the bird of prey could, in theory, engage a prey faster due cut a DNA sequence at defined locations or sites. The CRISPR/
to the associated decrease in aerodynamic drag. While the issue Cas9 method can be used to remove or disrupt specific genes,
of aerodynamic drag does not affect Hawkeye, the approach which is why it is popularly referred to as the genetic disruptor
could be of interest to Falcon, another member of the Aveng- (26), or, after a cut is made in the DNA, a new gene can be
ers. Falcon wears a flight pack that deploys wings enabling inserted at the location of the cut. CRISPR/Cas has been used
flight. Similar to a bird of prey, Falcon also needs to move his to study the development of fins in zebrafish (39), prevent the
head to observe targets, which could in turn lead to undesired browning of mushrooms (58), store images and films in living
fluctuations in the aerodynamic drag force, and possibly the lift bacteria (52), and remove the genetic code for muscular dys-
force, he experiences during flight. The hydrodynamic forces trophy from human embryos (30). The study on human em-
experienced by a real falcon have been investigated in the past bryos has implications for the modification of the human
by placing a live subject in a wind tunnel, where it was noted germline (18), as CRISPR/Cas is not entirely accurate and can
that, as the air speed increased, the falcon decreased wing span produce “off-target cuts” in a genome (11).
and lift coefficient (56). In the future, it may prove beneficial In theory, CRISPR/Cas could be used to replace human TFs
to model bird-of-prey aerodynamics using hydrodynamic co- with those from birds of prey in an effort to replicate the

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96 HAWKEYE EYE

photoreceptor cell density of birds of prey and Hawkeye. This avoid many of the ethical issues associated with genetic engi-
process is known as transgenesis, and in the past it has been neering.
used to modify goats to produce spider silk in their milk (6).
There are a number of other human genes that would also have Pedagogical and Communication Model
to be replaced with avian genes to fully replicate avian vision
abilities, such as the genes associated with the development of In the past, I have used popular culture icons to motivate
two foveae and the corneal accommodation mechanism. This interest among audiences in biological and physical sciences at
represents a difficult biological problem that could in the future schools, universities, and public events. From my experience,
be feasible using an advanced implementation of a tool such as scientific and engineering concepts can be interwoven with
CRISPR/Cas. Nonetheless, future geneticists will only be able superhero characters such as Hawkeye to create a connection
to explore transgenesis involving avian and human DNA, with an audience and enable communication of difficult-to-
subject to stringent evaluation and scientific scrutiny (18). grasp concepts. Such an approach can be referred to as the
Rather than rely on genetic engineering, we could replicate “middle-ground hypothesis” (62). The idea to write this article
Hawkeye’s eyesight by adapting bionic lenses. Companies and the related chapter in my book Secrets of Superhero
such as Ocumetics (43) and Alphabet (or Google) (5) are Science (13) stemmed from the positive engagement of audi-
developing bionic lenses that could replace lenses that have ences with the material.
cataracts and provide a patient with 20/20 vision. In the case of In the classroom environment, popular culture can make the
the Ocumetics bionic lens, the damaged biological lens in the learning process more engaging, invigorate the content linked
eye is replaced via cataract surgery, and the bionic lens is to key learning objectives, and potentially address issues stu-
connected to the ciliary muscles of the eye. This will provide dents have with key concepts (16). There are many ways that
the wearer with a larger range of vision capabilities that could a character like Hawkeye can be used in conjunction with
in theory be significantly greater than 20/20 vision. Alphabet’s learning objectives. First, teachers can focus on differences
bionic lens (patent pending) works in a slightly different way between standard human eyes and Hawkeye’s eyes to explain
from the Ocumetics lens. The Alphabet lens would be injected the function of the human eye and highlight limitations. Com-
into the eye and be integrated with the biological lens, if it is parison of human vision with that of birds of prey can also be
still present. The optical power of the lens can be varied using used to emphasize differences in the sensory systems of dif-
an external controller, and the lens could also communicate ferent animals. Second, the possibility of replicating the eye-
with external devices. These lenses are not designed to enhance sight of birds of prey in humans can prompt discussions on
a person’s eyesight beyond the limit of human vision. DNA, protein synthesis, advanced genetic engineering tech-
In addition there have been major advancements related to niques, such as the CRISPR/Cas9 genetic engineering tool, and
electronic retinas (38) and the stimulation of ciliary muscle transgenetics. These topics can, in turn, motivate dialogue with
response (17). Diseases such as retinitis pigmentosa and age- regard to responsible innovation and proper ethical practice in
related macular degeneration can have detrimental effects on genetics. Third, it is also possible to examine recent advance-
the photoreceptor cells in the retina and, as a result, lead to ments in modern technologies, such as wearable bionic lens,
blindness. In recent years, electronic retinal implants have been that can give the wearer enhanced eyesight or electronic
developed and implanted in patients in an effort to restore retinas. Fourth, Hawkeye can inspire student projects. Students
photoreceptor functionality (38). One example is the Argus II could investigate the limitations of Hawkeye’s sight and pres-
electronic epiretinal device, developed by the company Second ent strategies for how he might defeat villains under specific
Sight Medical Products, which has been shown to help a circumstances.
number of blind subjects identify letters and words (9). How- Hawkeye can form the basis for in-class assignments and
ever, an electronic retina such as the Argus II is a long way final examination questions. Relevant questions that could be
from fully reversing the effects of blindness caused by retinitis used in the classroom include the following:
pigmentosa or age-related macular degeneration. Nonetheless, 1. With reference to the The Avengers (2), discuss the
when the technology does reach this level, it offers the poten- limitations, if any, of Hawkeye’s eyesight as portrayed in
tial to be adapted for multiple advanced electronic foveae in the the final battle scene in New York City.
human eye, like those possessed by Hawkeye. In addition, in a 2. If one of Hawkeye’s eyes were damaged in a fight scene,
recent study, the ciliary muscles of 27 patients with early how would the functionality of his two foveae change?
emmetropic presbyopia were subjected to electrostimulation in 3. With reference to other muscle fibers in the human body,
an attempt to improve their near-vision capabilities (17). Re- are there any fibers in the body that could be used to
sults from the study showed that electrostimulation of the replace the ciliary muscles, such that Hawkeye’s lens
ciliary muscles did improve the vision of the patients. In the could be changed into shapes that are ordinarily inacces-
future this could prove to be a valuable course of action when sible?
treating the condition. On the other hand, electrostimulation of 4. If Hawkeye is able to see in the UV spectrum, would this
the ciliary muscles could be used to facilitate greater flexibility have any effect on his ability to target objects in the
and range of motion, thus facilitating larger deformation of the visible spectrum?
lens. In conjunction with the bionic lens mentioned earlier and 5. If we could use a genetic engineering tool such as
the possibility of electronic foveae, these treatments certainly CRISPR/Cas to modify human DNA to create an avian-
do open up the possibility of a technological route toward human hybrid, consider the different aspects of the eye
creating Hawkeye in real life. This certainly would be an easier that would have to be changed to duplicate bird of prey
path towards creating Hawkeye’s vision in the real world and vision.

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HAWKEYE EYE 97

In this paper I have motivated the discussion of Hawkeye birds of prey. Hawkeye can also motivate discussion in relation
with reference to one principal scene from the film The Aveng- to ligaments and tendons, bodily reflexes, and neural networks.
ers (2). However, there are many film scenes where Hawkeye He can also be used to discuss the ethical aspects of genetic
uses his advanced vision that could be used to motivate a engineering and allow students to reflect on the emergence and
scientific-based discussion. For example in Captain America: morality of emerging genetic engineering approaches. In addi-
Civil War (4), Hawkeye frees Captain America from Spider- tion, Hawkeye can be used to invoke interest in the latest
Man’s spider web by firing an arrow across an airport terminal. advancements in modern wearable technology, such as bionic
The arrow is seen passing straight through the spider web bound lens, and the implications of such technologies for future health
around the Captain’s hands. Based on the vision limitations of care.
birds of prey, an interesting exploratory study would to be esti- Outside of an educational environment, Hawkeye can be
mate how far away Hawkeye must have been when he fired the used to bridge the gap between academia and a nonspecialist
arrow to cut precisely through a small point in the spider silk. One audience. E. Paul Zehr has noted that “Pop culture contains
way of estimating the distance would be to use the average speed many compelling stories, and it makes sense to use these for
of an arrow and count the seconds from the time of release to the our purposes in communicating science” (62). While Hawkeye
time that it cuts through the arrow. However, would hawklike may not have the ability to fly like Superman or the healing
vision allow Hawkeye to resolve a small target only millimeters in powers of Wolverine, his powers and abilities fascinate many
size in the spider silk from that distance? people. He can be used as part of focused classroom teaching
For the general public, superheroes have a “wow” factor. or in the dissemination of new and innovative research. With-
Before the release of any superhero film, there is always out a doubt, Hawkeye can truly be a superhero for educators
increased media coverage where actors, the director, and film and scientific communicators.
studios seek to promote their latest production. Marketing of
these films uses TV advertisements, sharing of social media ACKNOWLEDGMENTS
content, and promotional posters on billboards and at bus stops. I thank Jean Philippe Frimat for stimulating discussions on the subject,
While some people may have never seen a superhero film, such sharing my enthusiasm for superheroes, and contributing an artistic image to
publicity campaigns ensure that most people are at least famil- the article.
iar with the superheroes. At the outset of most presentations or
DISCLOSURES
workshops, I ask the audience if they have seen a superhero
film. Almost everyone will have seen at least one superhero No conflicts of interest, financial or otherwise, are declared by the author.
film, while those that have not seen a superhero film are at least
AUTHOR CONTRIBUTIONS
familiar with characters such as Batman, Superman, Iron Man,
Spider-Man, and Wonder Woman, mainly due to the superhero B.F. conceived and designed research, performed experiments, analyzed
data, interpreted results of experiments, prepared figures, drafted manuscript,
film revolution. While the comic books can also be used to edited and revised manuscript, approved final version of manuscript.
communicate difficult scientific concepts (24), I have opted to
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