Mark D. Lenhart, and Walter J. Freeman
ABSTRACT: A physiological explanation of the long-term memory effect, Retroactive Interference, is proposed. We consider associative memory to be a dynamical system of cortical spatial patterns, manifested in the electrical fields of dendritic currents measured via electroencephalographic electrodes. Stimulus input causes state transitions with convergence toward a template pattern. In associative memories, new pattern storage affects all previously stored patterns. We believe RI is caused by accumulation of these changes, leading to failure of the system to converge toward a prior template pattern. Recordings from rabbit visual, somatic, and auditory cortices demonstrate pattern evolution over time with concomitant imposed learning. SUMMARY: Retroactive Interference, (RI) also called retrograde inhibition, is a well studied psychological phenomenon in both humans and animals. Typically studies have shown that recall performance for memories previously stored are negatively affected by later learning. Many hypothesis for this phenomena have been suggested. Among those commonly cited are memory address retrieval failure, memory trace decay, and competition. Scientists since Aristotle have been attempting to model forgetting in an effort to understand memory and the brain. Aristotle believed that memories undergo a passive decay over time. This hypothesis was embraced by E. L. Thorndike in 1914, who proposed a 'law of disuse' as the definitive factor of trace decay theory. It was the German scholar Hermann Ebbinghaus who first described Retroactive Interference in 1878. He created nonsense syllables and then methodically applied them to the study of memory recall performance. Using sequentially memorized lists of the syllables, he showed that forgetting could be attributed to the interfering effects of subsequently learned matter. His basic finding has been widely repeated and substantiated with a wide variety of human and animal subjects, learning and testing conditions. One predominant explanation for this finding is the two-factor theory of forgetting, which proposes that inhibition is due to both competition and unlearning. Specifically, forgetting is attributed to competition among memory traces during recall, and to unlearning of the original memory traces due to the subsequent learning. The unlearning hypothesis is equivalent to the experimental extinction found in animal studies. These and many modern cognitive psychology theories employ analogous or metaphoric structures, such as digital computers, to model the psychophysical data. It is unlikely the brain uses such computational algorithms. We are proposing another mechanism by which retroactive interference may occur. Findings derived from many years of research into the nature of perception in mammals has lead us to a new view of memory. We base our hypothesis upon empirical observations of overt natural behavior and correlated real-time electrophysiological events. We have found that cortical spatial amplitude patterns of a common carrier wave are the conveyor of perceptual memories and stimulus meaning. Associative memories in the mammalian brain are carried by globally distributed, chaotic, self organized dynamic activity patterns. Information retrieval is accomplished by external stimulus input causing the neural assembly to converge by state transition toward a specific, identifiable template pattern. This pattern is internally generated, and not directly dependent upon the external stimulus input pattern. It is profoundly influenced by the collection of past experiences of a particular animal. Memories stored as global attractor patterns manifest primarily internally generated stimuli meaning, rather than strictly transduced sensory data. Learning arises via Hebbian style synaptic strength changes influenced by correlated pattern output. Each learned class is a constraining of the dynamics of the cortical pattern into a stable, reproducible wing. Novel wings are formed by alterations of intracortical synapses during learning. Simultaneous excitation between neurons in the nerve cell assemblies are where new associations are integrated with all past associations. Three Albino New Zealand female adult rabbits were chronically implanted with sixty four channel (8x8 with inter-electrode distance of .8mm) EEG surface electrode arrays. The implant sites included the Somatosensory, Visual, and Auditory cortex. Computers recorded 6000ms of broadband (1-100 Hertz) electroencephalographic waves from familiarized animals in an awake, immobile state. No stimulus was delivered during recordings, however classical conditioning was performed before and midway through the duration of the experiment. Each recording session was held twice a week, with 22 total trials. Spectral decomposition, temporal filtering, and multivariate analysis of EEG segments were used to extract common wave form spatial amplitude patterns. Euclidean distance measures are calculated for the root mean square (RMS) carrier wave patterns generated from each weekly trial and used for tracking the evolution of carrier wave spatial amplitude patterns over time and between trials. The purpose of the study was to examine how electrophysiological (EEG) amplitude patterns evolve over time and with learning. A variety of factors can and do influence both the pattern and its rate of change. Several studies by Freeman have shown that in addition to the particular stimuli causing a subject to generate a specific pattern, the meaning of the stimuli, the environments it is given in, and the arousal state of the animal, can all have a profound influence upon the pattern. Our findings showed patterns typically mutate 5-15% from their starting points over the time span of a week. Prior to learning being imposed, the average intertrial change was 10%. The conditioning stimulus modality was changed for each animal after the 9th trial. The average percentile change between trial patterns where novel training was imposed jumped to 32% These findings are in concert with the theory that brains function with interconnected and distributed neural networks. In this associative view all new memory networks should have a small but noticeable effect on all previous memory stores. Our experimental findings of the evolution of EEG amplitude patterns are complementary to this theoretical framework. Forgetting and RI phenomena can be attributed to a difficulty in achieving or replicating previously shaped global cortical spatiotemporal patterns. The difficulty arises due to several factors. As new associative memories are formed in associative Hebbian networks, all previously stored memories are subtlety altered by the new connections. The original information is not written over, lost, or in competition with each other, but rather it is slowly modified over time, as the landscape of a mountain range is altered by weather and erosion over time. The exact point it becomes something new is not clear, and it will always retain some of its original character. In an associative memory, changes in any previously established memory, stored as spatial pattern, will have a small but significant impact upon all of the previously stored patterns. This characteristic is what allows an organism to bring all previous experiences to bear in analyzing a given stimulus input. Forgetting or retrieval failure or error can be caused by accumulation of these pattern changes due to extensive learning over time. These changes may lead to failure of the system to converge toward the correct template pattern. Experiments performed on three cortical implant sites in rabbits (visual, olfactory, and auditory) have provided data showing that these memory carrying patterns gradually evolve over time, and the rate of pattern change is influenced by imposed learning. References [1] Baird, B., Nonlinear dynamics of pattern formation and pattern recognition in the rabbit olfactory bulb, Physica D. 22, 150-77 (1986) [2] Freeman, W. J., Mass action in the nervous system, (1975) [3] Freeman, W. J.; Schneider, W., Changes in Spatial Patterns of Rabbit Olfactory EEG with Conditioning to Odors, 19 (1), 44 (1982) [4] Freeman, W. J.; Characterization of State Transitions in Spatially Distributed, Chaotic, Nonlinear, Dynamical Systems in Cerebral Cortex, 29 (3), 294-306 (1994)