Methods
Subjects and apparatus
Ten subjects, six women and four men, aged from 22 to 52 years, voluntarily participated in this study. All subjects were right-handed by self-report. Two of them were authors. The others were unaware of the purpose of the study. All participants had normal or corrected to normal vision. The experiment was part of a programme that has been approved by the local ethics committee.
Movements of the index finger and thumb were measured at a sampling rate of 250 Hz (resolution 0.1 mm) with an Optotrak 3020 infrared tracking system with two cameras. A small rigid body with three infrared markers (see Fig. 1) was fixed to the nail of the index finger and another one was fixed to the nail of the thumb of the subject’s right hand using an elastic gum.
The two objects were a black cylinder, made of polyoxymethylene (14 cm height, 4.5 cm diameter, 323 g mass, with a protrusion on its bottom), and a colourful opaque glass sphere (4.5 cm diameter, 123 g mass). A wooden board (60 × 52 cm) was placed on a height-adjustable table, onto which one of the objects was placed at one of nine locations on each trial. The locations formed a 3 × 3 grid with 10 cm spacing. The locations were marked by small indentations so that the objects could easily be placed at the correct positions, and the sphere would not roll away. Two starting positions were also marked: one centred near the subject (“near”) and one to the right of the grid (“side”). A schematic top view of the set-up can be seen in Fig. 2.
Procedure
Subjects stood in front of the table and its height was adjusted, if necessary, so that the task could be executed comfortably. Before each trial, subjects placed their right hand comfortably at one of the two possible starting positions, with the tips of the thumb and index finger touching each other. The experimenter indicated the starting position for each trial and placed the object at the appropriate location. He then gave a verbal signal to indicate that the subjects could start moving. Subjects were asked to grasp the object with a precision grip (holding it between index finger and thumb), to lift it and put it back at the same location, and then to move their hand back to the starting position. They could do so at their own pace. No further instructions or practice trials were given. A total of 36 conditions (2 starting positions × 2 object types × 9 object locations) were each presented once in each of five consecutive blocks, resulting in a total of 180 trials for each subject. Within each block, the conditions were presented in a random order.
Data analysis
Prior to data collection, a calibration trial was done in which the subject held an infrared marker between thumb and index finger. This marker’s position was later related to the positions of the sets of three markers on the rigid bodies attached to the fingernails. This allowed us to calculate the position in space of the part of the finger at which the hand-held marker had been, from the measured positions of the markers on the rigid body. The trajectory of that part of each finger was considered to be the fingertip’s trajectory towards the target object. Together, the two trajectories will be referred to as hand paths.
The speed of the hand was calculated by numerical differentiation of the average of the two digits’ positions. Movement onset was defined as the first frame in which the speed of the hand was higher than 50 mm/s. The moment of the grasp was defined as the last minimum of the mean of the two digits’ heights before this mean height reached its highest point (which always occurred when lifting the object). Movement time was defined as the time between movement onset and the moment of the grasp. Maximal grip aperture was defined as the maximal distance between the tips of the thumb and index finger, during the interval between movement onset and the moment of the grasp. We were mainly interested in the selected grasping points on the object, which can be expressed as the grip orientation at the moment of the grasp. Final horizontal grip orientation was defined as the orientation of the line connecting the two fingertips at the moment of the grasp, when projected onto the horizontal plane (Fig. 2). Final vertical grip orientation was defined as the angle between this horizontal projection and the line connecting the two fingertips. Horizontal grip orientation half way through the movement was calculated by determining the horizontal orientation on each trial at half the movement time.
In order to give an impression of the grasping movements, we determined average hand paths for each condition. The average positions were calculated by re-sampling the data of each digit’s path to give 50 equal steps (each step corresponding to 2% of the total path length) using linear interpolation between samples. Each of the 51 coordinates was then averaged across trials.
We were interested in whether differences in hand paths would affect the choice of grasping points. We expected the largest differences between the paths towards the two objects at the near right location. In order to quantify the extent to which the hand paths were different for the two objects when they were placed at this location, we took a straight line connecting the starting and grasping points of each digit and found the maximal deviation of each digit’s path from this line. We did so separately for each trial and separately for the horizontal and vertical plane. We averaged the maximal deviations across the five repetitions and compared these averages across objects and starting positions (for each digit’s starting position and plane). We also determined the grip orientation half way through the movement when moving towards the near right location, averaged the values across repetitions, and compared these averages across objects and starting positions.
The main dependent variables of our study were movement time, maximal grip aperture and final grip orientation. These values were determined for each trial and averaged across the five repetitions for each subject and condition. The standard deviation within the five repetitions was also determined and averaged across conditions and subjects to give a measure of the variability between movements. Effects on the average dependent variables were evaluated using 9 (object locations) × 2 (object types) × 2 (starting positions) repeated measures analyses of variance.
Besides these analyses, we also conducted 2 (object types) × 2 (starting positions) repeated measures analyses of variance for the grip orientation halfway through the movement and for the maximal vertical and horizontal deviations from the straight line connecting the starting with the grasping point of each digit. For these analyses, we only considered the trials when the object was placed at the near right location. For the deviation from the straight line, separate tests were done for the index finger and thumb.
All significant effects (P < 0.05) are mentioned in the results section. All significant interactions for the main dependent variables are illustrated with figures. Since we expected gradual systematic changes across object locations, we considered a graphical representation to be more informative than comparing effects at individual pairs of locations.
Results
Figure 3 shows an overview of the average grasping trajectories. The cylinder is grasped much higher than the sphere (Fig. 3b). For most object locations, the horizontal components of the digits’ paths to the spheres (continuous lines in Fig. 3a) are similar to those to the cylinders (dashed lines). However, for the objects at the near right location (as seen from above) they are not. When grasping the sphere at this location from the “near” starting position, the index finger moves over the object to reach the other side. When grasping the sphere at this location starting from the “side”, the thumb moves over the object (as illustrated in Fig. 1). When grasping the cylinder at this location from the “near” starting position, the index finger moves around the object to reach the other side. When grasping the cylinder at this location, starting from the “side”, the thumb moves around the object. Thus, the paths are much more curved in the horizontal plane when grasping the cylinder (Fig. 4). A similar but weaker tendency can be seen at other locations. At the location at which we expected the largest effect (the near right location), the difference between the maximal horizontal deviations for the two objects is indeed significant (F
(1,9) = 35.16, P < 0.001, for the index finger; F
(1,9) = 28.49, P < 0.001, for the thumb). There was no significant difference between the maximal vertical deviations.
Grip orientation half way through the movement to objects at the near right location also depends on the object type and starting position. There is a significant effect of starting position (F
(1,9) = 43.17, P < 0.001; on average 11.9 degrees further clockwise when starting from the “near” position) as well as a significant interaction between starting position and object type (F
(1,9) = 51.17, P < 0.001; the hand was oriented 12 degrees further clockwise when reaching for the cylinder when starting from the “near” position but 5 degrees further clockwise when reaching for the sphere when starting from the “side”).
Despite the differences in hand paths towards the objects, the final grip orientation is very similar for both of them (no main effect of object type; Fig. 3a). Final grip orientation depends on the starting position (F
(1,9) = 28.21, P < 0.001; on average it was 7.6 degrees further counter-clockwise when starting from the side). There is also a significant effect of object location (F
(8,72) = 203.18, P < 0.001; see Fig. 5a). There is a significant interaction between starting position and object location (F
(8,72) = 17.13, P < 0.001; Fig. 5a), and between starting position and object type (F
(1,9) = 6.23, P = 0.034; inset in Fig. 5a).
To summarize, final grip orientation appears to depend on more factors than only object location. The influence of object type is negligible: subjects grasp the sphere 0.9 degrees more counter-clockwise than they do the cylinder. There is no significant interaction between starting position, object location and object type, indicating that there is no difference between object locations with regard to the critical interaction between starting position and object type. The average standard deviation within replications for the final horizontal grip orientation is 4.6 degrees.
Movement time depends on the location of the target (F
(8,72) = 28.76, P < 0.001); it increases as a function of the distance from the starting position. A significant starting position by object location interaction (F
(8,72) = 12.34, P = 0.001) is consistent with this (Fig. 5b). There is no main effect of object type or starting position. However, there is a significant object type by object location interaction (F
(8,72) = 3.61, P = 0.001). This is mainly due to the fact that for objects at the near right location, movement times towards the cylinder were larger than the ones towards the sphere (Fig. 5c). The average standard deviation of the movement time across replications is 80 ms.
Maximal grip aperture depends on object type (F
(1,9) = 104.04, P < 0.001). The average maximal grip aperture when reaching for a cylinder is 7.1 cm, whereas for a sphere it is only 6.2 cm. This difference is not caused by the vertical grip orientation as the average angle that the grasping axis makes with the horizontal plane is close to zero (3 ± 6 degrees and 2 ± 7 degrees for the cylinder and the sphere, respectively, which cannot account for more than 1 mm of the larger grip when grasping the cylinder than when grasping the sphere). Object location also affects maximal grip aperture (F
(8,72) = 7.66, P < 0.001): the aperture is largest for object locations near the hand. There is a significant starting position by object location interaction (F
(8,72) = 3.66, P = 0.001; Fig. 4d). The average standard deviation of the maximal grip aperture across replications is 0.49 cm.
Discussion
When the objects were placed at the locations near the hand, subjects moved along different hand paths towards the two objects. The hand curved around the cylinder and moved over the sphere to grasp it. Movement time and maximum grip aperture were also affected. Nevertheless, subjects did not choose different grasping points. Despite the different hand paths, the final grip orientation was the same for the two objects.
It is likely that the difference in object height is responsible for the different hand paths. However, the objects also differed in shape and material. It is known that friction (Burstedt et al. 1999) and fragility (Gorniak et al. 2010; Savelsbergh et al. 1996), which are related to the material of which the object is made, can affect prehension, as can the shape of the object (Cuijpers et al. 2004). In order to test whether object height was really responsible for the differences in hand paths in experiment 1, a second experiment was carried out using cylindrical objects of different heights but of the same material.