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Precision Grips, Hand Morphology, and Tools
Precision Grips, Hand Morphology, and Tools
Precision Grips, Hand Morphology, and Tools
ABSTRACT This study asks whether there are discernable links between
precision gripping, tool behaviors,1 and hand morphology in modern homi-
noids, which may guide functional interpretation of early hominid hand
morphology. Findings from a three-pronged investigation answer this ques-
tion in the affirmative, as follows. (1) Experimental manufacture of early
prehistoric tools provides evidence of connections between distinctive human
precision grips and effective tool making. (A connection is not found between
the ‘‘fine’’ thumb/index finger pad precision grip and early tool making.)
(2) Manipulative behavior studies of chimpanzees, hamadryas baboons, and
humans show that human precision grips are distinguished by the greater
force with which objects may be secured by the thumb and fingers of one
hand (precision pinching) and the ability to adjust the orientation of gripped
objects through movements at joints distal to the wrist (precision handling).
(3) Morphological studies reveal eight features distinctive of modern humans
which facilitate use of these grips. Among these features are substantially
larger moment arms for intrinsic muscles that stabilize the proximal thumb
joints. Examination of evidence for these reveals that three of the eight
features occur in Australopithecus afarensis, but limited thumb mobility
would have compromised tool making. Also, Olduvai hand morphology
strongly suggests a capacity for stone tool making. However, functional and
behavioral implications of Sterkfontein and Swartkrans hand morphology are
less clear. At present, no single skeletal feature can be safely relied upon as an
indicator of distinctively human capabilities for precision gripping or tool making
in fossil hominids. Am J Phys Anthropol 102:91–110, 1997. r 1997 Wiley-Liss, Inc.
Functional analysis of fossil hominid 1986; Shrewsbury and Sonek, 1986; Brain et
hands generally includes an assessment of al., 1988; Ricklan, 1990). This ability is
their ability to use modern human precision considered a significant element of the evolv-
grips (Napier, 1959, 1961, 1962a,b, 1993; ing hominid hand/brain/tool complex. Infer-
Lewis, 1977, 1989; Susman and Creel, 1979; ences drawn from fossil hand bones about
Susman 1988a,b, 1989, 1991, 1994; Tuttle, the potential for precision gripping have
1981; Marzke, 1983; Marzke and Shackley, served sometimes to establish whether or
not a fossil hand was capable of both tool
using and tool making (Napier, 1959; Sus-
1The term ‘‘tool behavior’’ has been variously used in the
literature, in some cases implying exclusively tool making distinc- man, 1991). This paper points to flaws in
tive of humans (Susman, 1991) and in others referring variably such inferences from the Olduvai and Swart-
to tool using and/or tool-making abilities, some shared with us by
other animals (Susman, 1988a,b, 1994). In this paper the term is krans fossil hand bones, and proposes the
used to include both tool using and tool making behaviors of
humans and non-humans; the term ‘‘tool making’’ is used in place
of ‘‘tool behavior’’ whenever the discussion is focused upon
distinguishing a capacity for removing flakes from stone pre- Received 9 May 1994; Accepted 2 October 1996.
forms from a more general capacity to manipulate stone tools. *Correspondence to: Mary W. Marzke.
use of eight morphological features to pre- way between radial and ulnar deviation.
dict precision grip capabilities that facilitate This classic paper simplified the analysis of
habitual, effective prehistoric stone tool mak- hand postures and movements by demon-
ing. Evidence in the literature to support the strating that these fall into two categories,
use of these features rests on investigation one (precision grips) involving the thumb
involving (1) the experimental replication of opposed to the fingers and allowing preci-
prehistoric tools by archaeologists, (2) the sion control of objects, and the other (power
comparison of three catarrhine species in grips) incorporating the palm of the hand as
manipulative behavior, and (3) the biome- well, to secure the object as an extension of
chanical analysis of hand and wrist morphol- the forearm for use in forceful activities such
ogy in chimpanzees. as hammering.
The investigation asks the following: What The ‘‘precision’’ term is often restricted to
precision grips are essential to habitual and grips in which a small object is held between
effective manipulation of prehistoric stone the thumb’s distal phalanx and the terminal
tools? Are these grips shared with other volar pads of one or more fingers. This is the
catarrhine species? Are there morphological type of grip specified or implied in discus-
features distinctive of each species which sions of the potential for precision gripping
are essential to the effective use of their
by fossil hominid species in several papers
respective precision grip repertoires? Is there
by Napier (1959, 1961, 1962a,b, 1964),
fossil evidence for the origin and evolution of
Krantz (1960), Susman (1979, 1988a,b, 1989,
morphological features that specifically re-
1991), and Susman and Creel (1979). My
flect precision grip capabilities, including
definition is considerably broader. A focus on
human precision grip capabilities that facili-
tate effective prehistoric tool making? The the thumb pad/distal index pad grip re-
answers are drawn by integrating my col- stricts the amount of information on preci-
laborative research results with those of sion gripping and tool-making potential that
other investigators. The primary goal of the might be gleaned from the structure of fossil
paper is to bring into focus the full, inte- hominid hand bones. Kinematic and electro-
grated body of evidence for morphological myographic studies of the human hand take
adaptations to prehistoric tool making, which account of a much greater variety of thumb/
cannot easily be discerned in reports of the finger postures and movements used in se-
separate research projects. curing, holding, and maneuvering objects
with precision (see, for example, Landsmeer,
HISTORICAL REVIEW: THE PRECISION 1962; Long et al., 1970; Long, 1981; Chao et
GRIP QUESTION al., 1976, 1989). In particular, the impor-
tance of the ability to maneuver, as well as to
The term ‘‘precision grip’’ is defined here
retrieve and hold objects with precision,
as any grip that involves the thumb and one
seems to have been almost completely ig-
or more fingers, with or without the palm
nored by physical anthropologists, with the
serving passively as a prop. It is distin-
exception of Krantz (1960). Therefore, I have
guished from the term ‘‘power grip,’’ in which
objects are strongly squeezed by the fingers adopted a definition of precision gripping
alone or squeezed by the fingers, thumb, and that is more reflective of what the hand
actively by the palm. The first term was actually does in tool use.
introduced by Napier (1956), who defined it Over time the discussion of precision grips
as one in which an object is pinched between became reduced in the physical anthropologi-
part or the whole flexor aspect of the fingers cal literature to the claim that the potential
and the opposing thumb. He related the grip for one type of precision grip (between the
to morphological facilities for abduction and distal thumb and distal index finger pad) is
medial rotation at the carpometacarpal and linked to one aspect of hand morphology
metacarpophalangeal thumb joints, flexion (relative thumb length) and to a capacity for
and abduction at the finger metacarpopha- stone tool making. In my view these links
langeal joints, and wrist dorsiflexion mid- have not been established; no simple test in
PRECISION GRIPS, HAND MORPHOLOGY, AND TOOLS 93
hand morphology exists for establishing tool- toric stone tools by humans; (2) the system-
making as opposed to exclusively tool-using atic comparative study of precision gripping
potential in fossil hominids. In particular, in humans and other catarrhine species; and
Susman’s (1988a,b, 1989) arguments from (3) the biomechanical analysis of hands in
the morphology of Paranthropus to precision humans and nonhuman species.
gripping, and from precision gripping to tool
making are not supported by the criteria on Precision grips actually used in the
which he relies, which include (1) relative manufacture of prehistoric stone tools
thumb length (Susman, 1988a) and (2) links
How does one perform the experiments
between Olduvai hand morphology, thumb–
and make the evaluations in the absence of
fingertip precision gripping, and tool mak-
living representatives of prehistoric hominids?
ing (Susman, 1988b). Evidence regarding (1)
Three indirect approaches provide clues to
is not provided for Paranthropus, and Na-
the minimal requirements of hand morphol-
pier (1962a) largely dismissed (2) because he
ogy for effective precision manipulation of
could not find evidence for relative thumb
the tools, and to features that enhance the
length in the Olduvai hand, and he was able
behaviors. These are (1) the experimental
(like Krantz, 1960) to reproduce early homi-
replication of prehistoric stone tools by ar-
nid stone tools without using the precision
chaeologists (Marzke and Shackley, 1986),
grip. It is regrettable that the focus on
(2) observation of stone tool making by con-
hominid hand evolution has become almost
temporary groups that still use stone tools
exclusively concentrated on a single type of
(Toth et al., 1992), and (3) the experimental
precision gripping behavior, about which
replication of prehistoric stone tools by our
insufficient evidence is available from the
close pongid relatives (Schick and Toth, 1993;
fossil record, and whose importance for Old-
Toth et al., 1993).
owan tool making is not established. A larger
body of evidence is evaluated below for Experiments in the manufacture of stone
possible links between morphology, preci- tools. Marzke and Shackley (1986) found
sion gripping, and tool behaviors. A new grip that the replication and use of Oldowan
classification scheme is presented, which tools elicited primarily three grips: the pad-
reflects this broader approach and is de- to-side grip between the thumb and the side
signed to facilitate a comprehensive test for of the index finger, the 3-jaw chuck (‘‘base-
both tool using and tool making, as more of ball’’) grip, and the cradle precision pinch
the fossil record is revealed. grip by the thumb and four finger pads (see
Figs. 1 and 2). Strong pinch as well as
RECENT APPROACHES
precision handling of stones by the thumb
Since humans can make stone tools with pad and the side of the index finger (with
an ape-like power grip, should we abandon and without buttressing by the other fin-
the hypothesis that the human hand evolved gers) were effective for cutting with flakes
in adaptation to tool behaviors, and the against strong resistance, and also (with
expectation that fossil hominid hand mor- buttressing) for stabilizing stone preforms
phology will reflect stages in the evolution of in one hand against blows by a hammer-
tool behaviors? Rather than rejecting the stone in the other. The 3-jaw chuck pinch
notion, we need to abandon untested as- grip with precision handling is the most
sumptions of links between hand morphol- effective one for throwing, and for the strong
ogy, precision gripping, and tool making, pinch and precision maneuvering of round
and to test whether the hominid hand hammerstones when they are used to re-
evolved in adaptation to economical and move flakes from stone cores. The cradle
effective manipulation of tools. Currently pinch and handling grip occurred frequently
three areas of investigation are providing when firm pinch and maneuvering of large
insights into possible links between several stone preforms by the thumb and four fin-
types of precision gripping, hand morphol- gers of one hand were required during flake
ogy, and effective tool behaviors. These are: removal by hammerstones. The grip in-
(1) the experimental replication of prehis- volves extensive movement of the thumb in
94 M.W. MARZKE
Fig. 1. Human precision finger grips. A: Two-jaw tip-to-tip. B: Two-jaw pad-to-pad. C: Two-jaw
pad-to-side. D: Three-jaw full finger pad-to-pad (‘‘baseball’’).
opposition to all four fingers. All these grips handling allowed for rapid core reduction. If
provided the extensive contact surfaces on a thumbless power grip of the core is used,
the hand needed to control large preforms or the core must be repositioned for each strike
hammerstones by one hand alone. The facil- either by dropping it and retrieving it in a
ity for generating a strong precision pinch new orientation or by shifting its position
provides the ability to expose a greater with the hand holding the hammerstone.
surface of a stone than would be possible if There is savings in time and effort by preci-
the stones were embedded in the palm and sion handling.
fingers in power grips. The exposed surface These three grips have not extensively
can be struck by the hammerstone or used in been discussed in connection with fossil
pounding and digging without injuring the hominids. The 2-jaw pad-to-pad precision
fingers. hold of small objects was not recruited in
The same grips were again prominent these activities. Nor were power grips, which
during a recent experiment in which hand actively incorporate the palm. This latter
muscle recruitment during stone tool use observation seems curious in the light of
and tool making was monitored with electro- experiments in stone tool making by Krantz
myography, during simultaneous videotap- (1960) and Napier (1962b, 1964), in which
ing of the hands (Marzke et al., abstract they showed that some tools could be made
submitted). Especially striking was the con- using the power grip exclusively. However,
stant repositioning of the stone core by the the purpose of their experiments was to
thumb and fingers of the non-dominant hand, determine whether these tools could be made
in preparation for each strike of the hammer- without a well-developed human thumb,
stone by the dominant hand. This precision because the authors thought at the time
PRECISION GRIPS, HAND MORPHOLOGY, AND TOOLS 95
that early hominid morphology facilitated to the efficient manufacture of stone tools
only ape-like power grips. This was similar comparable to those found in the prehistoric
to asking whether this paragraph could be record. By observing archaeologists engaged
typed by the fifth finger. Certainly the poten- in the manufacture of prehistoric tools we
tial is there, but the paragraph is more can evaluate the grips which have proved to
effectively generated, with less expended be most effective in providing control, preci-
energy and less repetitive stress on a single sion, and a minimum of injury and stress on
joint, by ten well-trained fingers including the bones and joints. We found (Marzke and
the thumb. Our experiments with tool manu- Shackley, 1986; Marzke et al., abstract sub-
facture suggest that stone tool use is so mitted) that only the three precision grips
greatly facilitated by capabilities for apply- listed above (along with precision handling),
ing firm precision pinch grips as opposed to and no power grips, consistently expedite
power grips, that only with the former capa- tool use and tool making with relatively
bility would tool making have become ha- little muscle fatigue and discomfort in the
bitual. Thus the results of the Napier and hand joints. This strongly implies that cer-
Krantz experiments were interesting, but tain distinctive human features of the hand
not in the end relevant. might reasonably be explained as adapta-
My collaborative work has led me to view tions to increasing demands for these preci-
the evolution of the human hand from a sion grips with the evolution of prehistoric
different perspective. Enormous advantages tools.
to tool use and tool making are provided by An objection to this line of reasoning has
modern human hand morphology. From this been raised by Sarmiento (1994), who states
vantage point, we may determine which that ‘‘Because tools are fashioned to fit the
elements of our grip repertoire, with their hand that makes them, it is difficult, if not
morphological underpinnings, are essential illogical, to interpret the anatomical struc-
96 M.W. MARZKE
ture of the human hand as adaptive to tool using a hammerstone. This finding further
use.’’ We disagree. For at least 2.5 million highlights the importance of focusing on
years the irregularly shaped raw materials forceful, one-handed manipulation of stones
used to fashion tools were held and wielded in tracing the morphological basis of homi-
directly by hominid hands. The varied uses nid tool making.
of these raw materials stressed the joints of
the hands, and could not have been executed Precision gripping in chimpanzees and
habitually and expeditiously unless the joints hamadryas baboons
increasingly possessed ranges of movement, Prehistoric stone tool manipulation does
and constraints on mobility that assured not necessarily elicit the much-discussed
effective grips with minimal repetitive stress thumb pad-to-index pad grip (Marzke and
and joint loading. We argue that our hand Shackley, 1986), but instead involves a num-
must have evolved first in adaptation to ber of precision grips that are both dynamic
using these raw materials effectively, then and forceful, and are not frequently de-
to making tools and to using them habitu- scribed in the literature on human and
ally for a variety of manipulative activities nonhuman manipulative behavior. We initi-
that would have been impossible without ated observations of manipulative behavior
evolutionary adaptations in bone and joint in several nonhuman primate species. Cir-
morphology to the stresses involved. cumstances under which precision grips are
elicited were recorded, grips were catego-
Contemporary stone tool-making behav-
rized according to the positions of the thumb
ior. Videotapes of hafted stone axe manu-
and fingers, and it was noted whether the
facture by a New Guinea group studied by grips involve maneuvering of objects by the
Toth et al. (1992) reveal the same emphasis thumb and fingers, or forceful holding of
as our experiments, on cradle, 3-jaw chuck, objects against resistance.
and pad-to-side grips. They also reveal an
absence of thumb pad/index finger pad grips, Chimpanzees. Marzke and Wullstein
during the removal of flakes from the stone (1996) observed five precision grips used by
axe preforms. It is not until the stage at captive chimpanzees to retrieve and/or hold
which the axes are hafted by the New Guinea foods, tools, and other objects of various
tool-makers, by pinching fiber with the sizes by the thumb and fingers (Table 1). The
thumb and finger pads and winding it around most common of these were (1) a pad-to-side
the axe and its handle, that the latter grip is hold between the thumb and the side of the
recruited. index finger and (2) a cup hold, in which an
object is supported passively by the up-
Nonhuman primate stone tool-making turned palm and propped by one or more
behavior. A bonobo (‘‘Kanzi’’), trained to fingers, held in parallel, and by the thumb,
remove flakes from stone cores with a ham- held at varying angles of opposition to the
merstone, has demonstrated that human- fingers. (This grip is distinguished from
like hands are not necessary for this activity power grips by the lack of enclosure and
(Schick and Toth, 1993; Toth et al., 1993). squeezing of the object against the palm.)
However, he developed another technique The chimpanzees also used holding grips
for flake removal on his own which has between the thumb tip (not the pad) and the
become his favorite (Schick and Toth, 1993). tip, distal pad, or dorsal aspect of a finger,
The technique is to throw stones forcefully for retrieving small objects (for example, the
with one hand against a hard tile floor, or tip/tip hold during grooming). A grip be-
against a cobble on the ground. It achieves tween the full thumb and index finger pads
flake removal without requiring control of a was not observed. The pad-to-side grip was
stone by one hand against external forces. used for holding fruit rinds during feeding,
Toth et al. (1993) found that the impact for stripping leaves from stems, and for
forces between the stones were much larger one-handed probing with or holding of ob-
in throwing against the cobble than those jects against moderate resistance. When
the chimpanzee was capable of producing maintenance of the pad-to-side or cup hold
PRECISION GRIPS, HAND MORPHOLOGY, AND TOOLS 97
TABLE 1. Chimpanzee precision grips used for retrieving and holding food and other ob-
retrieval and holding of objects
jects against mild resistance, but two hands
Number of (using the pad-to-side grip) were called into
fingers Thumb/finger Thumb/finger/
(‘‘jaws’’) in grip position palm position play when a more firm hold was required, as
in breaking hard biscuits, for example. The
Precision Precision fin-
finger grips ger/passive thumb and index finger were not observed to
palm grips precisely maneuver prehended objects.
2-jaw chuck Tip-to-tip hold
(thumb/index) Pad-to-tip hold The pad-to-pad hold has also been ob-
Pad-to-dorsal served by Rose (1977) in olive baboons feed-
hold
Pad-to-side hold
ing in the wild and by Jolly (1970) in Theropi-
3-5-jaw chuck Cup hold (thumb/ thecus gelada. Napier (1961) noted that
(Thumb/2–4 palmar sur- terrestrial cercopithecines in general closely
fingers) faces of fingers;
object rests approach humans in the capacity for this
against grip.
upturned
metacarpals)
Contrasts in precision gripping between
humans and the two nonhuman species.
Modern humans exhibit all the precision
was threatened by external forces, such as grips described above for retrieving and
pulling of citrus fruit pulp from the rind by holding of small objects by chimpanzees and
the teeth, the other hand (and occasionally a hamadryas baboons. They also use the pinch
foot as well) was recruited. It appears that and handling grips listed in Table 2 and
the thumb and fingers together are not able illustrated in Figures 1 and 2. The human
to generate a firm enough balanced pinch to precision gripping repertoire appears to be
resist more than moderate forces dislodging distinctive in three ways. First, humans
the object. This finding is consistent with often apply considerable force to these grips,
the behavior of the bonobo ‘‘Kanzi’’ during
achieving precision pinch as opposed to less
tool making. The chimpanzee thumb and
firm precision holding. Landsmeer (1962)
fingers also do not precisely maneuver ob-
and others stress the delicacy of precision
jects; changes in the orientation of an object
grips, but many of our modern tools also
are effected primarily by movements at the
require a firm pinch (without a power
wrist and forearm joints, or by transferring
the object to the lips and retrieving it again squeeze by the palm) to meet resistance, as,
by the hand in the desired orientation.2 A for example, in our experimental stone tool
distinction is therefore made in the grip making. Second, humans accommodate the
classifications of Tables 1 and 2, between the thumb and fingers to the shape of objects,
precision holding grips of chimpanzees and abducting and rotating the fingers to bring
the precision pinch and handling grips of their volar pads into contact with the sur-
humans. face of the object, further enhancing security
of the grip. This is seen, for example, in the
Hamadryas baboons. Guthrie (1991) and 3-jaw chuck grip of hammerstones by the
Jude (1993) reported for hamadryas ba- archaeologists in our experiments. Third,
boons the pad-to-side and pad tip-to-pad tip humans adjust the orientation of the object
holding grips described above for the chim- (i.e., they translate and rotate the object)
panzee as well as a pad-to-pad hold, in through movements at the carpometacar-
which both the thumb and index distal pal, metacarpophalangeal and interphalan-
interphalangeal joints are extended to bring geal joints of the thumb and fingers, as
the volar pads over the distal phalanges into described by Landsmeer’s (1962) term ‘‘pre-
opposition. These grips were effective for cision handling.’’ These movements are fine-
tuned by displacements in the cushion-like
2These grips are also described and/or illustrated in recent pads on the distal phalanges, and facilitate
studies by Christel (1993) and Boesch and Boesch (1993), which activities such as the rapid reorientation of
approach chimpanzee manipulative behavior from other perspec-
tives and are discussed in Marzke and Wullstein (1996). stone cores for flaking by the archaeologists.
98 M.W. MARZKE
TABLE 2. Human precision grips for retrieval, holding, pinching, and handling of objects
Number of
fingers in grip Thumb/finger position Thumb/finger movements
Precision finger pinch grips Precision handling
2-jaw chuck Tip-to-tip Tip/pad translation and pad/pad rotation
(thumb/index Pad-to-pad
finger) Pad-to-tip
Pad-to-side Translation and rotation
3-jaw chuck Tip-to-tip Tip/pad translation and pad/pad rotation
(thumb/index/ Distal finger pad-to-pad
medius) Full finger pad-to-pad Pad/pad rotation and translation
form a pattern which is favorable to achiev- 33% in hylobatids and 24% in chimpan-
ing the firm precision pinch grips and preci- zees and orangutans. Results of a recent
sion handling that in our experiments (Mar- comparative kinematic analysis of thumb
zke and Shackley, 1996; Marzke et al., muscles in a sample of 8 chimpanzees
abstract submitted) appear to be essential to and in 2 samples of 7 humans each
effective tool making. The features are de- (Marzke et al., 1995; Marzke et al., in
scribed below, and our recent findings are press) show that three of these muscles
incorporated under number 3. have a much greater potential in humans
1. Broad ungual tufts. Susman (1979) has for stabilizing the thumb during strong
shown that humans have proportionately pinch grips of objects. Applying tech-
broader tufts on the distal phalanges niques used at the Mayo Clinic for mea-
than apes. The tufts support pads whose suring muscle force potential and tendon
large surfaces distribute pressure during moment arms (Chao et al., 1989; An et
forceful grasping and whose mobility al- al., 1991; Linscheid et al., 1991; Horii et
lows accommodation of the pads to un- al., 1993), it was found that the human
even surfaces as well as fine-tuning in oblique adductor pollicis and opponens
the positioning of objects. The difference pollicis muscles have not only larger
is illustrated in Shrewsbury and Johnson physiological cross-sectional areas (PCSA,
(1983, Fig. 1), who confirm Susman’s a measure of potential force) but also
observation. The large friction surface larger tendon moment arms (MA) for
would have been essential for securing flexion/extension at the carpometacarpal
and controlling the cradle pinch of large joint than are found in the chimpanzee
preforms and the 3-jaw grip of hammer- sample (Table 3). It can be seen from
stones in habitual tool making. Table 3 that the potential torque (PT, a
2. A long thumb relative to the length of the product of PCSA and MA) is therefore
fingers. Humans are distinctive among approximately 15 times that of chimpan-
primates in having the longest thumb zees for the oblique adductor and almost
relative to the second finger (Napier, triple the chimpanzee PT for the oppo-
1993). This is well illustrated in Schultz nens. Also at the carpometacarpal joint
(1969). The distal pad of the thumb is the human flexor pollicis brevis MA for
therefore able to control objects of varied abduction/adduction is 4.7 times that of
sizes and shapes against the volar as- the chimpanzee sample. Since the PCSA
pects of extended or moderately flexed is the same for the two species, humans
fingers during pinch and translation and have 4.7 times the PT in flexor pollicis
rotation of the objects. A long thumb is brevis because of the difference in MA
important to prehistoric tool making be-
(Table 3). When human pinch grips con-
cause it can control stones with the volar
centrate opposing force at the ends of the
aspect of extended or only partially flexed
digits, very large moments of stabilizing
fingers on the opposite side of the stone.
muscles are required at the proximal
The longer the thumb relative to the
fingers, the greater the control of the joints (Brand and Hollister, 1992). Thus,
stone. Fully modern human thumb/finger during the manufacture of stone tools,
proportions probably would have been the more forceful the hammering, the
essential only for tasks requiring firm more muscle torque is required to stabi-
pinch and precision handling of small lize the proximal thumb joints in main-
tools by the thumb and the full pads of taining the pad-to-side, 3-jaw chuck, and
hyperextended distal phalanges. cradle pinch grips of the stones. Our
3. Proportionately well-developed intrinsic kinematic findings have pinpointed the
muscles of the thumb. Tuttle (1969) found joint regions and muscle attachment ar-
that the thumb intrinsic musculature eas whose comparative biomechanical
constitutes 39% of total intrinsic hand analysis will clarify details of joint func-
musculature in humans, compared with tion relevant to the precision grips most
100 M.W. MARZKE
TABLE 3. Intrinsic thumb muscle mean physiological cross-sectional areas, tendon moment arms, and torque
potential at the trapeziometacarpal joint, exhibiting marked differences between chimpanzee and human samples1
PCSA MA PT
Muscle Activity Species (cm2 ) (cm) (cm3 )
Adductor pollicis (oblique head) Carpometacarpal flexion/ext. Human 3.13 2.35 7.36
Chimpanzee 1.25 0.38 0.48
Opponens pollicis Carpometacarpal flexion/ext. Human 2.63 1.25 3.29
Chimpanzee 1.50 0.74 1.11
Flexor pollicis brevis (superficial head) Carpometacarpal abduction./add. Human 1.42 0.98 1.39
Chimpanzee 1.42 0.21 0.30
Metacarpophalangeal flexion/ext. Human 1.42 0.63 0.89
Chimpanzee 1.42 0.40 0.57
Adductor pollicis (transverse head) Carpometacarpal abduction/add. Human 0.89 1.66 1.48
Chimpanzee 1.25 3.32 4.15
1 Chimpanzee data are from a recent study by Marzke et al. (in press). The human moment arms are from a just-completed study at
Mayo Clinic, Rochester, MN, kindly provided prior to publication by P. Smutz. Human PCSA data are from Linscheid et al. (1991).
PCSA: physiological cross sectional area; MA: moment arm; PT: potential torque (PCSA 3 MA).
Fig. 4. Joints between the second metacarpal, trapezoid, and trapezium. A: Dorsal view of chimpanzee
(left) and human (right). B: Volar view of chimpanzee (left) and human (right). Note in the human the
volar-proximal orientation of the metacarpal joint surface for the trapezium (arrow).
a roughened volar surface on the ungual area of the proximal compartment. While
tuft reflecting the attachments of fascial the features associated with compartmen-
fibers which restrict mobility of this com- talization may not be essential to firm
partment, thus reducing the amount of precision grip and precision handling of
pinch pressure necessary to maintain stones, they probably would have en-
hold of objects (Shrewsbury and Johnson, hanced these behaviors, since deforma-
1983; see especially Figs. 1, 2, 4, and 5). tion of the compartmentalized pads al-
Comparative dissections and analyses of lows finely controlled translation and
grips utilizing this region of the thumb rotation of stones. Their occurrence in
and fingers by Shrewsbury and Sonek fossils, together with other features essen-
(1986) indicate that this compartmental- tial to the behaviors, would be indicative
ization of the tufts into a mobile proximal of potentially very effective precision ma-
and a less mobile distal section is unique nipulation by the thumb and distal finger
to humans, and that it permits a variety pads.
of precision postures (tip-to-tip, pad-to-
pad, and pad-to-tip). These are capable of Additional morphology claimed to be
exploiting individually or in combination distinctive of humans. Susman (1994)
the firmness of pinch afforded by the discussed another feature, a broad first meta-
distal compartment together with the carpal head relative to metacarpal length,
cushioning, mobility, and larger sensory which he considers to be distinctive of hu-
PRECISION GRIPS, HAND MORPHOLOGY, AND TOOLS 103
Fig. 5. Distal surface of the right capitate in a Chimpanzees. We were surprised to dis-
chimpanzee (left) and human (right). The facet for the cover a set of three morphological features
third metacarpal occupies the entire surface in the in chimpanzees that appear to be adapta-
chimpanzee, with the second metacarpal surface at
approximately right angles to it on the radial side. In the tions to their manipulative behavior, in a
human, the concave second metacarpal surface may be hand which is otherwise quite highly special-
seen to the right of the third metacarpal surface, facing
more distally than in the chimpanzee. ized in its long fingers and strong finger
flexor musculature for suspensory behavior
(Marzke et al., 1994). The features, which
mans. He attributes the greater relative enhance their pad-to-side holding grip, in-
breadth in humans to the presence of three clude the following. First, a saddle surface
muscles (flexor pollicis longus, flexor pollicis for the metacarpal on the trapezium which
brevis deep head, and a first palmar interos- is narrow and convex in its anteroposterior
seus) that are lacking in chimpanzees, and dimension, particularly on its ulnar aspect
suggests that human tool behavior explains (Guthrie, 1991), so that during opposition of
the larger number of muscles. This reason- the thumb the proximal volar beak on the
ing is then applied to the functional inter- metacarpal rides up on the trapezial convex
pretation of metacarpal head size in fossil saddle, locking against it. This locking mech-
hominid species; nonhuman manipulative anism is capable of stabilizing the joint
behaviors are inferred from a narrow head during pressure of the thumb tip against
in A. afarensis and human-like tool behav- objects held along the side or volar aspect of
iors are inferred from a broader head in the the index finger, but it limits excursion of
Swartkrans fossils. the metacarpal in opposition to the fourth
These hypothesized links between joint and fifth fingers. Second, a distal interpha-
surface breadth, muscle number, and tool langeal joint of the thumb which generally
behavior are open to serious question. First, cannot fully extend (Tuttle, 1969, 1970;
the joint surface breadth was compared only Shrewsbury and Sonek, 1986; Marzke and
in humans and chimpanzees. Hamrick and Wullstein, 1996). The thumb is thus posi-
Inouye (1995) and Ohman et al. (1995) ap- tioned in such a way that its tip is directed
plied Susman’s measurement to gorilla toward the index finger when an object is
thumb metacarpals, which proved to fall held between them. In humans, the flexor
within the human range of variation. Sec- pollicis longus muscle, which is frequently
ond, the number of muscles is not in itself absent in chimpanzees (Tuttle, 1970; Mar-
relevant to a comparative analysis of joint zke, 1971), is a dynamic positioner of the
surface size relative to muscle function. What distal phalanx. Third, the adductor pollicis
matters is the total amount of torque gener- muscle sends a tendon to the distal phalanx
ated by muscles at the metacarpophalan- of the thumb. This tendon appears to func-
geal joint. Third, the three human muscles tion as a constraint on extension of the distal
104 M.W. MARZKE
phalanx. (This extra tendon has been ob- muscle at the level of the wrist inserts into
served in 16 of 18 chimpanzee cadavers in the base of the distal phalanx of the thumb,
which the insertion was specifically exam- in a position to effect the application of
ined during dissection in my laboratory, as pressure by the volar pad against the object
well as in gorilla and orangutan specimens. held on the volar pad of the index finger.
It has also been reported by Tuttle, 1969, These findings on hamadryas baboons re-
1970.) In addition, the transverse portion of turn the thumb pad/index finger pad grip
the adductor pollicis muscle has approxi- from the exclusive realm of ‘‘refined’’ human
mately three times the potential torque (PT) behavior to membership among grips merely
of the human muscle for thumb abduction/ shared with humans. Morphological corre-
adduction at the trapeziometacarpal joint lates of the grip in the baboons should be
(Table 3; Marzke et al., in press). considered among features that might have
The significance of our behavioral/morpho- existed in the hominid ancestral hand prior
logical findings in chimpanzees to the inves- to abandonment of quadrupedal locomotion.
tigation of prehistoric hominid manipula-
tive behavior is that they reveal both a large EVOLUTION OF THE HUMAN HAND,
repertoire of effective precision grips for PRECISION GRIPPING, AND TOOL
retrieving and holding objects, with morpho- MAKING
logical correlates for one of them, and limita-
In this section we consider how the infor-
tions on the forcefulness and maneuverabil-
mation in Recent Approaches may be ap-
ity of these grips. These findings provide
plied to the functional and behavioral inter-
new morphological clues to possible manipu-
pretation of fossil hominid hand morphology.
lative adaptations in early fossil hominid
hands, but they also put into relief the
Predictions from the comparative
distinctive aspects of human precision grip-
hominoid studies
ping whose morphological correlates should
also be sought in fossil hands. The human morphological features de-
scribed in Recent Approaches are significant
Hamadryas baboons. The morphological in their combined ability to facilitate firm
features of particular interest here are those pinch and precision handling of stones by
facilitating the thumb pad-index finger dis- each hand alone. This ability is likely to
tal pad grip, the so-called ‘‘acme’’ of human have become essential to hominids when
precision gripping stressed in the literature they grew increasingly dependent upon tools
on hominid hand evolution. The features are manufactured by the removal of flakes from
built into a morphological pattern special- core preforms. It is predicted that the fea-
ized (with relatively short phalanges) for tures will appear in the fossil record when
semi-digitigrade locomotion. First, the index the prehistoric record indicates a trend from
finger is particularly short relative to the opportunistic flake production toward ha-
thumb (Etter, 1973). Second, the distal inter- bitual, widespread and systematic Oldowan
phalangeal joints are capable of extending tool making. One would then expect to find
fully and hyperextending, allowing much of skeletal evidence for the following: (1) thumb
the distal volar pad of the thumb to oppose sufficiently long relative to finger length to
its counterpart on the index finger (Jude, permit pinch and maneuvering of stone cores,
1993). This pad-to-pad grip thus provides a hammerstones, and flakes, (2) intrinsic
relatively large surface area for maintaining muscle torque potential sufficient to main-
hold of small objects. Third, as in the chim- tain the pinch of these stones against strong
panzee and humans, there is a saddle joint resistance, (3) metacarpophalangeal joint
with incongruent surfaces (Rose, 1992) be- surfaces allowing some degree of cupping by
tween the trapezium and first metacarpal, the fingers, and (4) broad distal phalanges
which allows the thumb pad to be brought which could distribute the load associated
into opposition to the pad of the index finger. with strong pinch grips. None of these fea-
Fourth, a tendon originating from the cen- tures in isolation would be sufficient evi-
tral fibers of the flexor digitorum profundus dence for tool making.
PRECISION GRIPS, HAND MORPHOLOGY, AND TOOLS 105
Note that it is likely that most of the least the stage 1 capacities seen in chimpan-
morphological features that facilitate firm zees for tool behavior. There is a set of
precision pinch grips, and perhaps even morphological features in the chimpanzee
precision handling, evolved before the ad- thumb which are best explained by their
vent of stone tool making with increasing frequent use of a pad-to-side precision hold-
dependence upon the use of unmodified ing grip (Marzke and Wullstein, 1996). The
stone, bone, and wood tools. The effective- presence of these features in very early fossil
ness of activities such as probing, digging, hominid hand bones would suggest a similar
nut-cracking, cutting, and throwing with precision holding specialization, and would
these tools would have increased with the predict potential for behaviors that exploit
ability to orient the tools precisely and to this gripping capability in chimpanzees.
hold them firmly in one hand against resis-
tance. Therefore, caution is urged in attempt- Stage 2: Hadar. Australopithecus afaren-
ing to distinguish an advanced stage of tool sis has three of the eight human features
using from an early stage of tool-making described in Recent Approaches, which
capabilities among fossil hands. Presence of should have facilitated two distinctively hu-
these features in fossil species permits confi- man precision grips (together with precision
dence in their potential for habitual, effective handling): the firm pad-to-side pinch and
tool making, but the features do not necessarily the 3-jaw chuck pinch. One of the human
confirm that the species were tool-makers. features (no. 6) is a cam-like projection on
the volar radial side of the second metacar-
Stages in the evolution of tool behavior pal head and dorsal beveling of the articular
What constitutes evidence of adaptations surface (personal observation; Fig. 3), which
to tool behavior in fossil hominid hands? I would have caused pronation of the index
define four stages through which tool behav- finger as it flexed. This capacity was comple-
ior may have progressed. Stage 1 consists of mented by feature 7, which is orientation of
occasional use and modification of natural the second metacarpal joints with the capi-
objects for tools, with moderate control by tate and trapezium away from the sagittal
hands specialized primarily for quadrupedal plane. This orientation reflects a capacity
locomotor demands. This stage is illustrated for slight ‘‘give’’ of the metacarpal base in
by living chimpanzees, whose tool behaviors pronation. The features together allow for
in the wild have been described by McGrew rotation of objects by the thumb and side of
(1992) and by Boesch and Boesch (1993). the index finger and for positioning of the
With stage 2, there would have been increas- full volar surface of the index finger on
ing use of natural objects of various sizes spherical objects to secure a 3-jaw chuck
and shapes as tools, facilitated by new, hold grip. The human orientation of the
distinctively human morphologic features carpometacarpal joints also provides for ac-
which increased pinch strength, finger con- commodation of axial load, indicating that
trol, and tolerance of new stresses associ- the joints may have been regularly sustain-
ated with grips of the tools. Stage 3 mani- ing strong index finger muscle contraction
fests a dependence upon the manufacture as associated with a firm pad-to-side pinch of
well as controlled use of unhafted stone objects. The third feature is a longer thumb
tools. Stage 4 is characterized by the manu- relative to the fingers than in chimpanzees
facture and use of hafted tools and very (no. 2; Marzke, 1983). This should have
small tools, some of which probably required provided an advantage in securing and con-
controlled pad-to-pad pinch by a thumb and trolling flakes with the middle and distal
index finger of human proportions. segments of the flexed fingers in the pad-to-
side grip, and in holding relatively large
Fossil evidence for stages in the evolution stones by the three-jaw chuck grip. The
of precision grip capabilities and tool thumb probably was not long enough to
behaviors facilitate pad-to-pad precision pinch and han-
Stage 1. It is likely that the ancestor we dling of small tools by the thumb and distal
share with African apes would have had at finger pads. Cradle precision gripping by the
106 M.W. MARZKE
thumb and all four fingers of one hand may features associated with other precision
have been enhanced by a facility for supina- grips. A distal thumb phalanx attributed to
tion at the fifth metacarpophalangeal joint. Paranthropus is described by Susman
The fifth metacarpal head has an asymme- (1988b) as similar to that of modern humans
try that mirrors the pattern of the second in breadth (no. 1), reflecting a large overly-
metacarpal head (Fig. 3) and would have ing pad that would have facilitated firm
guided the proximal phalanx into supina- pinch of objects against pressure by the
tion with flexion. However, control of objects fingers. It is also described as similar in the
by this grip may have been limited by trape- presence of an insertion area for the flexor
ziometacarpal morphology. The base of the pollicis longus muscle (no. 4). Susman is
thumb has a chimpanzee-like projecting vo- silent on the presence or absence of features
lar first metacarpal beak, which probably that allow compartmentalization of the pulp
stabilized pad-to-side grips but interfered on the broad distal phalanx (no. 8). He
with smooth excursion of the metacarpal in argues that a broad proximal joint surface,
opposition to the fourth and fifth fingers, stout shape, and marked crest for attach-
where it is needed to control large objects ment of the opponens pollicis muscle on a
with the finger pads (Marzke, 1992). first metacarpal (SKX 5020), along with the
Tool using by the thumb, index and third broad pollical distal phalanx with an exten-
fingers would have been enhanced by the sive fossa marking the insertion of the long
pad-to-side and 3-jaw chuck precision capa- flexor muscle, reflect a capacity for a preci-
bilities, but tool making would have been sion grip comparable to that of modern
restricted by the limited potential for a firm humans (Susman, 1988b). None of these
precision cradle grip and precision handling features, with the exception of pollical distal
with this grip. Pad-to-side pinch and preci- phalangeal breadth, has yet received the
kind of systematic comparative analysis ap-
sion handling may have been used to apply
plied by Susman and by others to the 8
pressure to tools in activities such as cutting
morphological features on which the present
and probing or digging. The capacity for
discussion is based.
holding stones in a 3-jaw chuck grip would
There is no evidence in the available bones
have been an advantage for stone-throwing
of morphology that might have limited 3-jaw
(Marzke, 1983).
chuck and cradle grip capabilities necessary for
Stage 2: Sterkfontein. The fossils from tool making. Information about morphological
this site have three of the human features features that would confirm a potential for
listed in Recent Approaches, which would these grips and behaviors is unavailable.
have facilitated at least the human pad-to- A second thumb metacarpal (SK 84) has a
side precision pinch and handling grips. marked volar projection of the articular
Ricklan (1987, 1990) describes a distal thumb surface on the head (Napier, 1959). Napier
phalanx (STW 294) which is broad (no. 1), notes that the degree of projection is greater
with a tuft and spines reflecting compart- than he observed in his human and ape
mentalization of the pads (no. 8), and with sample. It is likely to have provided an
dimensions and a tendon insertion mark advantageous moment arm for metacarpo-
indicating a well-developed flexor pollicis phalangeal flexion/extension by the flexor
pollicis brevis muscle. Since the muscle in
longus muscle with greater mechanical ad-
humans has a larger moment arm compared
vantage than the human muscle. Informa-
with chimpanzees (Table 3), and plays an
tion on features 2 and 5–7 is not available at
important role in stabilizing pinch grips
present to indicate whether other precision
involving the distal finger pads, the morphol-
grips involving the finger pads could have
ogy of this Swartkrans metacarpal head has
exploited these features of the thumb.
implications for precision grip potential in
Stage 2: Swartkrans. Here human fea- the species to which it belonged.3
tures of the thumb are found (nos. 1 and 4),
3The SK 84 metacarpal has been attributed by Susman (1988b)
which are compatible with a firm pad-to-side to Homo erectus, rather than to Paranthropus, on the basis of a
grip, but information is not available about feature on the metacarpal head that he reports is shared with a
PRECISION GRIPS, HAND MORPHOLOGY, AND TOOLS 107
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