The vergence system is responsible for the convergence and divergence movements of the eyes. The first quantitative study of vergence dates to the work of Rashbass and Westheimer in
1961. Based on their experimental results, they modeled vergence as a simple feedback control system (Rashbass & Westheimer,
1961). However, more recent behavioral data support a dual control paradigm. Jones dissected the vergence response into a transient and sustaining component (Jones,
1980). The Dual-Mode Theory also describes vergence as a two-component system that includes a transient and a sustaining component (Hung, Semmlow, & Ciuffreda,
1986; Semmlow, Hung, & Ciuffreda,
1986) The transient portion is assumed to be an open-loop control component that enhances the speed of a vergence response. The sustaining portion is assumed to be driven by a visual feedback, closed-loop control system that provides fine tuning of the response to enable the extraordinary accuracy seen in binocular fixation.
Investigators often use the terms prediction and adaptation in different ways. Prediction is a higher level process evoked when a stimulus is in some way predictable. In the life sciences, classical adaptation usually describes a process that requires a significant training period to change a behavior based on environmental conditions and that persists with a lasting aftereffect. From an engineering viewpoint, adaptive control is simply a modification of a system parameter irrespective of the time course of that change or its aftereffect. In some cases, adaptive control can be closely related to prediction (Deno, Crandall, Sherman, & Keller,
1995).
Several investigations have studied oculomotor behavior in response to periodic stimuli where the timing, magnitude, and direction of the stimulus could be predicted. Prediction is typically marked by anticipatory movements as in the smooth pursuit system (Kowler,
1989), saccadic system (Kalesnykas & Hallet,
1987), and vergence (Alvarez, Semmlow, Yuan, & Munoz,
2002; Kumar, Han, Garbutt, & Leigh,
2002; Yuan, Semmlow, & Munoz,
2000b). In smooth pursuit, utilizing prediction led to a decrease in response latency and tracking error (Barnes & Asselman,
1991a; Barnes & Asselman,
1991b; Barnes, Barnes, & Chakraborti,
2000; Deno et al.,
1995; Kowler,
1989). Finally, prediction will substantially reduce response latency in both saccadic and vergence eye movements (Krishnan, Farazian, & Stark,
1973; Rashbass & Westheimer,
1961).
Adaptive behavior has also been observed in several oculomotor systems. The saccadic system can be manipulated utilizing a decreasing or an increasing gain protocol based on double step stimuli (Albano,
1996; Deubel,
1995; Semmlow, Gauthier, & Vercher,
1989; Straube & Deubel,
1995; Straube, Fuchs, Usher, & Robinson,
1997). The gain of the smooth pursuit system can also be modulated via adaptation (Churchland & Lisberger,
2002). The disparity vergence system also shows clear adaptive behavior: changes in the tonic vergence or phoria occurs in response to sustained demand (Patel & Firth,
2003; Schor, Gleason, Maxwell, & Lunn,
1993). Because this sustained demand is usually produced by prisms, this adaptive modification is termed “prism adaptation.”
Recent studies have shown that disparity vergence dynamics can also be modified, but as we show here, these changes maybe due to a type of prediction (Munoz, Semmlow, Yuan, & Alvarez,
1999). Munoz et al. (
1999) utilized an experimental protocol based on step-ramp stimuli (a step change in vergence immediately followed by a ramp) to increase the apparent gain of the system, resulting in convergence movements with large overshoots. Their findings indicate that the transient vergence component was strongly modified by these step-ramp stimuli. They also found a positive correlation between the maximum velocity of a subject's normal response and the increase in response velocity modified by the step-ramp conditioning stimuli (Munoz et al.,
1999). Another study by Takagi et al. (
2001) used a double-step protocol similar to that used in saccadic eye movements to both increase and decrease the gain. Studying only convergence eye movements, they showed that the peak velocity increased with an increasing gain protocol and peak velocity decreased with the decreasing gain protocol (Takagi et al.,
2001).
In the current study, four different experiments were performed: in three experiments, the stimulus direction, but not the timing, could be predicted; whereas in the fourth experiment, timing, direction, and amplitude could not be predicted. In all experiments, one or more “conditioning stimuli” were more prevalent than the stimulus used to “test” the modification. In the first three experiments, the dynamics of both convergence and divergence responses could be increased or decreased by conditioning using the appropriate repetitive step stimulus. One experiment showed that the dynamics of large responses could be decreased whereas the dynamics of small responses increased by the same intermediate conditioning stimulus. In the fourth experiment, when all predictive clues were eliminated, the absence of modification suggested that change in dynamics was the result of some type of prediction. Yet none of our stimuli were completely predictive because the stimulus onset was randomized. Moreover, the behavior observed was quite different from that seen in response to a truly predicable stimulus. Such stimuli, such as a periodic square wave, produce anticipatory eye movements and a decrease in latency, but not a change in movement dynamics. The modifications of this research show neither anticipatory movements nor latency changes but do show significant changes in movement dynamics.