Sparks (1989) argues, from converging behavioral and single-unit data in alert macaques, that saccade targets cannot be encoded by retinal error (RE) alone. The decisive experiment is the stimulate/compensate paradigm: on ~30% of trials, the superior colliculus was electrically stimulated after a briefly flashed target disappeared, driving the eyes to a new orbital position. Monkeys nonetheless made saccades to the actual target location in space—impossible under pure retinocentric control, and unpredictable enough that compensation could not be preprogrammed .
The review then maps three representations onto anatomy: (1) RE in retinotopic superficial SC, with receptive fields shifting with gaze; (2) motor error (ME) carried by quasi-visual (QV) cells in intermediate SC, which fire based on the required saccade vector whether created by a new target or by an eye movement after target disappearance; QV-like memory cells were concurrently found in the intraparietal cortex (Gnadt & Andersen); and (3) tentative head/body-centered signals in the thalamic internal medullary lamina and area 7a, where visual responses are multiplicatively gain-modulated by gaze angle—a population code from which head-centered position can be decoded (Zipser & Andersen) .
Figure: qualitative strength assigned from the review's own stated confidence—author notes head-centered mapping evidence is "not yet compelling."
As a single-author narrative review by the originator of the key paradigms, it carries an inherent confirmation-risk: the strongest evidence (stimulate/compensate) comes from Sparks' own lab, and cross-lab replication was limited at the time. The author commendably flags this by acknowledging early studies used "informal and uncontrolled test procedures" and that a topographic head-centered map had not been found . Additional blind spots from a modern vantage: the review is strictly macaque-based (human generalization untested); the distributed gain-modulation account leaves open whether "head-centered" is truly explicit or implicit; and later work (e.g., LIP recordings in marmosets and macaques showing heterogeneous visual/post-saccadic signals; transsaccadic sensory memory in V4/MT) has both extended and complicated the simple three-representation picture .
Bottom line: the core claim—saccade targeting uses spatial, not purely retinal, coordinates—is very well supported (behavioral + neurophysiological + causal stimulation). The claim that explicit head-centered maps exist is explicitly flagged by the author as unresolved. What would change the conclusion: direct evidence that compensatory saccades persist with corollary discharge pathways blocked (i.e., relying only on proprioception, which Guthrie et al.'s muscle-elimination control argues against), or demonstration that a purely retinal feed-forward model can reproduce double-step accuracy.
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