title: Behavioral variability, elimination of responses, and delay-of-reinforcement gradients in SHR and WKY rats.
abstract: BACKGROUND
  Attention-deficit/hyperactivity disorder (ADHD) is characterized by a pattern of inattention, hyperactivity, and impulsivity that is cross-situational, persistent, and produces social and academic impairment. Research has shown that reinforcement processes are altered in ADHD. The dynamic developmental theory has suggested that a steepened delay-of-reinforcement gradient and deficient extinction of behavior produce behavioral symptoms of ADHD and increased behavioral variability.
  
  
  METHODS
  The present study investigated behavioral variability and elimination of non-target responses during acquisition in an animal model of ADHD, the spontaneously hypertensive rat (SHR), using Wistar Kyoto (WKY) rats as controls. The study also aimed at providing a novel approach to measuring delay-of-reinforcement gradients in the SHR and the WKY strains. The animals were tested in a modified operant chamber presenting 20 response alternatives. Nose pokes in a target hole produced water according to fixed interval (FI) schedules of reinforcement, while nose pokes in the remaining 19 holes either had no consequences or produced a sound or a short flickering of the houselight. The stimulus-producing holes were included to test whether light and sound act as sensory reinforcers in SHR.Data from the first six sessions testing FI 1 s were used for calculation of the initial distribution of responses. Additionally, Euclidean distance (measured from the center of each hole to the center of the target hole) and entropy (a measure of variability) were also calculated.Delay-of-reinforcement gradients were calculated across sessions by dividing the fixed interval into epochs and determining how much reinforcement of responses in one epoch contributed to responding in the next interval.
  
  
  RESULTS
  Over the initial six sessions, behavior became clustered around the target hole. There was greater initial variability in SHR behavior, and slower elimination of inefficient responses compared to the WKY. There was little or no differential use of the stimulus-producing holes by either strain. For SHR, the reach of reinforcement (the delay-of-reinforcement gradient) was restricted to the preceding one second, whereas for WKY it extended about four times as far.
  
  
  CONCLUSIONS
  The present findings support previous studies showing increased behavioral variability in SHR relative to WKY controls. A possibly related phenomenon may be the slowed elimination of non-operant nose pokes in SHR observed in the present study. The findings provide support for a steepened delay-of-reinforcement gradient in SHR as suggested in the dynamic developmental theory of ADHD. Altered reinforcement processes characterized by a steeper and shorter delay-of-reinforcement gradient may define an ADHD endophenotype.
authors.affiliation: Department of Physiology, Institute of Basic Medical Sciences, University of Oslo, Oslo, Norway. e.b.johansen@medisin.uio.no.
authors.forename: Espen B
authors.initials: EB
authors.lastname: Johansen
authors.forename: Peter R
authors.initials: PR
authors.lastname: Killeen
authors.forename: Terje
authors.initials: T
authors.lastname: Sagvolden