Non-associative Form Research Articles

Clouding and Aggregation Behavior of PPO-PEO-PPO Triblock Copolymer (Pluronic®25R4) in Surfactant Additives Environment

Abstract Aqueous solutions of commercially available Pluronic®R i. e. triblock copolymer composed of polypropylene oxide(PPO)-polyethylene oxide(PEO)-polypropylene oxide(PPO), (code named 25R4, BASF) in the presence of surfactant additives viz. hydrotropes (NaTS, NaXS), anionics (SDS, SDBS) and cationics (DTAB, CTAB) were investigated by cloud point(CP), surface tension and conductance techniques. All the techniques clearly indicated that Pluronic®25R4 interacts strongly with anionic surfactants but weakly with cationic surfactants. Surfactant additives used here, produced a marked increase in the CP of the copolymer. Elevation in CP is attributed to the solubility of PPO block of the copolymer and formation of Pluronic-surfactant complex. The effect of NaCl salt on the CP of Pluronic®25R4 at added SDS and CTAB was also studied and it was decreased due to the screening effect. At the presence of 25R4 in its non-associated form, the surface tension and conductance data is discussed with binding process and indicated strong interaction of Pluronic®25R4 with SDS, showing two break points (P1 and P2) in the plot and very weak interaction with CTAB observing single break point.

During the stationary growth phase, Yarrowia lipolytica prevents the overproduction of reactive oxygen species by activating an uncoupled mitochondrial respiratory pathway

In the branched mitochondrial respiratory chain from Yarrowia lipolytica there are two alternative oxido-reductases that do not pump protons, namely an external type II NADH dehydrogenase (NDH2e) and the alternative oxidase (AOX). Direct electron transfer between these proteins is not coupled to ATP synthesis and should be avoided in most physiological conditions. However, under low energy-requiring conditions an uncoupled high rate of oxygen consumption would be beneficial, as it would prevent overproduction of reactive oxygen species (ROS). In mitochondria from high energy-requiring, logarithmic-growth phase cells, most NDH2e was associated to cytochrome c oxidase and electrons from NADH were channeled to the cytochromic pathway. In contrast, in the low energy requiring, late stationary-growth phase, complex IV concentration decreased, the cells overexpressed NDH2e and thus a large fraction of this enzyme was found in a non-associated form. Also, the NDH2e–AOX uncoupled pathway was activated and the state IV external NADH-dependent production of ROS decreased. Association/dissociation of NDH2e to/from complex IV is proposed to be the switch that channels electrons from external NADH to the coupled cytochrome pathway or allows them to reach an uncoupled, alternative, ΔΨ-independent pathway.

Hidden nonassociative structure in supersymmetric quantum mechanics

AbstractIt is shown that the Hamilton equations in supersymmetric quantum mechanics can be presented in nonassociative form, where the Hamiltonian is decomposed into two nonassociative factors.

Towards an image of a memory trace

Learning and memory lead to functional and structural changes in the brain, ultimately providing a basis for adaptive behavior. The honeybee is an elegant model for the study of learning and memory formation as it permits both the visualization of neural activity related to the events occurring in olfactory learning and the behavioral assessment of olfactory learning (Galizia and Menzel, 2000). The formation of odor memories in the honeybee is thought to involve the two primary processing centers of the olfactory system, the antennal lobe (AL) and the mushroom body (MB). The intrinsic neurons of the MB – the Kenyon cells (KCs), located within the lip region of the MB calyx – are the site of convergence of the neural pathways that transmit odor information from the projection neurons (PNs) of the AL and reward information from the VUMmx1 neuron (Hammer, 1997). In recent years, imaging studies performed in the honeybee AL and MB lip have indicated that pairing odor and reward induces changes in neural activity (Faber and Menzel, 2001; Faber et al., 1999), reinforcing the anatomical suggestion that KCs are likely to undergo associative plasticity during learning. This study, Szyszka et al. (2008) provides new evidence that odor-evoked activity of KCs can be modified by sensory experience, and the changes can be associative or non-associative. Szyszka et al. (2008) visualized odor representations in the MB of the honeybee by imaging the calcium responses of a subpopulation of KCs – the clawed KCs, by retrogradely labeling the cell population using dye injections into the ventral alpha/gamma lobe. This allowed the authors to image, in a relatively selective way, the bulk dendritic activity of KCs over neuropil. Szyszka et al. (2008) first examined KC activity in response to repeated stimulation with the same odor and found that it led to a decrease in KC responses, a non-associative form of plasticity resembling a phenomenon previously described in locust PNs (Stopfer and Laurent, 1999). Separate experiments found that honeybee PNs did not show repetition-induced depression, indicating that the effect is not a generalized run-down. The authors then compared KC activity before and after pairing one odor with sucrose reward and tested how the associative pairing procedure affects KC responses to the rewarded odor (CS+) and to the unrewarded odor (CS−). During pre-training, KC responses to both CS+ and CS− decreased with repeated stimulation. After pairing CS+ with sucrose CS+ responses recovered and CS− responses decreased further (Figure ​(Figure1).1). Intriguingly, examination of spatiotemporal patterns within the imaged region before and after conditioning suggested additional changes in both CS+ and CS− evoked KC activity following conditioning. Figure 1 KC responses are subject to associative and non-associative plasticity. (A) Scheme of honeybee antennal lobes and mushroom bodies (MB). Calcium responses in the somata and dendrites of clawed Kenyon cells (clawed KCs, red) were recorded in the MB calyx ... The results of this study support the suggestion that KC odor responses can be modified by sensory experience and that these changes can be associative, extending the earlier work of the Menzel group (Faber and Menzel, 2001; Faber et al., 1999). Although a causal link between this activity and learning and memory remains to be established, the strengthening of support for a leading candidate cellular substrate is an encouraging observation that can be followed up in several ways. Firstly, it should be possible to examine the same cell population at the molecular and cellular levels in order to elucidate the processes underlying these modifications. Secondly, the changes in spatiotemporal patterns hint at restructuring of KC odor representations that might reflect a correlate of information storage. It may now be feasible, using similar imaging approaches but at single cell resolution, to resolve the details of the changes. Finally, it should be possible, albeit technically challenging, to take advantage of the possibility of behavioral measurements in combination with imaging to try to directly link KC changes to memory storage itself.

Intermediate-Term Memory is Modulated by the Circadian Clock

Sensitization of the tail-siphon withdrawal reflex in Aplysia, a nonassociative form of learning, affords a superb opportunity to investigate the regulation of learning and memory by the circadian clock. The circadian clock has been shown to modulate long-term but not short-term sensitization. However, no previous studies have examined the role of the circadian clock in intermediate-term memory. Noxious stimulation delivered to the side of the animal using a spaced training protocol resulted in canonical intermediate-term sensitization dependent upon both MAPK signaling and protein synthesis. The authors found that intermediate-term sensitization exhibited strong rhythms in expression in both light-dark cycles and constant darkness. Animals trained during the (subjective) day demonstrated significantly more intermediate-term memory than animals trained at night. Baseline responses prior to training were not modulated by the circadian clock. Thus, these results indicate that the circadian clock strongly modulates intermediate as well as long-term memory.

Lasting changes in a network of interneurons after synapse regeneration and delayed recovery of sensitization

Regeneration of neuronal circuits cannot be successful without restoration of full function, including recovery of behavioral plasticity, which we have found is delayed after regeneration of specific synapses. Experiments were designed to measure neuronal changes that may underlie recovery of function. Sensitization of the leech withdrawal reflex is a non-associative form of learning that depends on the S-interneuron. Cutting an S-cell axon in Faivre’s nerve disrupted the capacity for sensitization. The S-cell axon regenerated its electrical synapse with its homologous cell after 3–4 weeks, but the capacity for sensitization was delayed for an additional 2–3 weeks. In the present experiments another form of non-associative conditioning, dishabituation, was also eliminated by S-cell axotomy; it returned following regeneration. Semi-intact preparations were made for behavioral studies, and chains of ganglia with some skin were used for intracellular recording and skin stimulation. In both preparations there was a similar time-course, during 6 weeks, of a lesion-induced decrease and delayed restoration of both S-cell action potential threshold to depolarizing pulses and S-cell firing in response to test stimuli. However, the ability of sensitizing stimuli to decrease S-cell threshold and enhance S-cell activity in response to test stimuli did not fully return after regeneration, indicating that there were lasting changes in the circuit extending beyond the period necessary for full recovery of behavior. Intracellular recordings from the axotomized S-cell revealed a shift in the usual balance of excitatory and inhibitory input, with inhibition enhanced. These results indicate that loss of behavioral plasticity of reflexive shortening following axotomy in the S-cell chain may be related to reduced S-cell activity, and that additional processes underlie full recovery of sensitization of the whole body shortening reflex.

Nitroxide Bound β-Cyclodextrin: Is There an Inclusion Complex?

236CDTIPNO, a derivative of the persistent free radical TIPNO (1,1-dimethylethyl 2-methyl-1-phenylpropyl nitroxide) covalently bound to a permethylated-beta-cyclodextrin, was prepared. Self-association of 236CDTIPNO in water was proved by solvent- and competition-dependent EPR spectroscopy experiments with 2,6-di-O-Me-beta-cyclodextrin (DIMEB) and permethylated-beta-cyclodextrin (TRIMEB) as external hosts competing for accommodation of the TIPNO moiety. Temperature-dependent EPR spectra were simulated with a novel two-dimensional (field-temperature) EPR simulation program that afforded a full determination of the thermodynamic parameters characterizing the rate constants of the self-inclusion reaction derived from Arrhenius and Eyring models. This method allows separating the line broadening effects due to relaxation from a chemical exchange, even if only the fast exchange regime is accessible experimentally. The activation parameters for the forward and backward steps were consistent with an equilibrium between a nonassociated form and a weakly associated form, with activation free enthalpies for each reaction of around 34 kJ.mol(-)(1).

Generalization of amygdala LTP and conditioned fear in the absence of presynaptic inhibition

Pavlovian fear conditioning, a simple form of associative learning, is thought to involve the induction of associative, NMDA receptor-dependent long-term potentiation (LTP) in the lateral amygdala. Using a combined genetic and electrophysiological approach, we show here that lack of a specific GABA(B) receptor subtype, GABA(B(1a,2)), unmasks a nonassociative, NMDA receptor-independent form of presynaptic LTP at cortico-amygdala afferents. Moreover, the level of presynaptic GABA(B(1a,2)) receptor activation, and hence the balance between associative and nonassociative forms of LTP, can be dynamically modulated by local inhibitory activity. At the behavioral level, genetic loss of GABA(B(1a)) results in a generalization of conditioned fear to nonconditioned stimuli. Our findings indicate that presynaptic inhibition through GABA(B(1a,2)) receptors serves as an activity-dependent constraint on the induction of homosynaptic plasticity, which may be important to prevent the generalization of conditioned fear.

Circadian modulation of complex learning in diurnal and nocturnal Aplysia.

Understanding modulation of memory, as well as the mechanisms underlying memory formation, has become a key issue in neuroscience research. Previously, we found that the formation of long-term, but not short-term, memory for a nonassociative form of learning, sensitization, was modulated by the circadian clock in the diurnal Aplysia californica. To define the scope of circadian modulation of memory, we examined an associative operant learning paradigm, learning that food is inedible (LFI). Significantly greater long-term memory of LFI occurred when A. californica were trained and tested during the subjective day, compared with animals trained and tested in the subjective night. In contrast, animals displayed similar levels of short-term memory for LFI when trained in either the subjective day or night. Circadian modulation of long-term memory for LFI was dependent on the time of training, rather than the time of testing. To broaden our investigation of circadian modulation of memory, we extended our studies to a nocturnal species, Aplysia fasciata. Contrary to the significant memory observed during the day with the diurnal A. californica, A. fasciata showed no long-term memory for LFI when trained during the day. However, A. fasciata demonstrated significant long-term memory when trained and tested during the night. Thus, the circadian clock modulates memory formation in phase with the animals' activity period. The results from our studies of circadian modulation of long-term sensitization and LFI suggest that circadian modulation of memory formation may be a general phenomenon with potentially widespread implications for many types of long-term learning.

Trauma and resulting sensitization effects are modulated by psychological factors

Traumas have both immediate consequences and proactive consequences. Examples include learned helplessness, HPA-axis responsivity, gastrointestinal vulnerability to ulcer, and other correlates of anxiety disorders. Both immediate and proactive consequences may be modulated by behavioral and cognitive evolutionary evolved adaption processes, among which are forms of learning that enable 'coping'. Examples of associative and non-associative forms of coping and effects on learned helplessness, HPA-axis responsivity, and gastrointestinal vulnerability are presented. The importance of attention to behavioral contingencies in situations in which potentially traumatic events occur is emphasized as critical to understanding that it is not the physical event(s) per se that determine the immediate and long term consequences.

Detection and purification of two antibody–antigen complexes via selective adsorption on lowly activated anion exchangers

Taken advantage of the mechanism of adsorption of macro-molecules on ionic exchangers, (a multipoint interaction between the protein and the support), it is possible to selectively adsorb large proteins leaving small ones in the supernatant. Associated proteins should present a significant difference in its size as compared to the non-associated forms. Thus, the protein complexes may have much larger surfaces to interact with the support. Here, by selecting the support with the highest activation degree that was unable to adsorb the non-associated proteins, we have shown the simple and selective adsorption of immuno complexes (as a model), while antibodies and antigens remained in the supernatant. Therefore, it was possible to selectively adsorb on lowly activated supports (e.g., agarose 4BCL having only 1μmol of amino groups per g of support) rabbit IgG/anti-rabbit immunoglobulins (immuno complex), while these supports were unable to adsorb the individual immunoglobulines. Similarly, horseradish peroxidase (HRP)/anti-HRP were selectively adsorbed on lowly activated supports, while the individual proteins were not adsorbed at all. Afterwards, the adsorbed associated proteins (purified at least from the non-associated counterparts and concentrated by the adsorption on the support) may be cross-linked with aldehyde-dextran and be desorbed from the matrix for their analysis. This strategy may permit very simple experiments to detect the presence of protein–protein complexes. Finally, we have shown the advantages of this technique compared to the use of one of the proteins previous immobilized on a support.

Nucleus accumbens dopamine receptors in the consolidation of spatial memory.

Nucleus accumbens dopamine is known to play an important role in motor activity and in behaviours governed by drugs and natural reinforcers, as well as in non-associative forms of learning. At the same time, activation of D1 and D2 dopamine receptors has been suggested to promote intracellular events related to neural plasticity. Therefore, in this study we wished to investigate the role of the two classes of dopamine receptors within the nucleus accumbens on the consolidation of spatial information. On day 1, CD1 male mice were placed in an open field containing five different objects and, immediately after three sessions of habituation, the animals were focally injected within the nucleus accumbens with either the D1 antagonist SCH 23390 (12.5, 25 or 50 ng/side), or the D2 antagonist sulpiride (25, 50, 75 or 100 ng/side). Twenty-four hours later the ability of mice to discriminate an object displacement was assessed. Both the D1 and the D2 antagonists impaired the ability of mice to detect the spatial change. If the highest doses of the two antagonists were injected 2 h after the end of the last of the habituation sessions, no effect was observed in the reactivity to spatial change examined 24 h later. These data demonstrate that activation of both D1 and D2 receptors within the accumbens is necessary in the early stages of the consolidation of spatial information. The data are discussed in terms of involvement of nucleus accumbens dopamine in information processing in the absence of explicit reinforcers.

Miscibility and phase behavior in thermosetting blends of polybenzoxazine and poly(ethylene oxide)

Thermosetting polymer blends composed of polybenzoxazine (PBA-a) and poly(ethylene oxide) (PEO) were prepared via in situ curing reaction of benzoxazine (BA-a) in the presence of PEO, which started from the initially homogeneous mixtures of BA-a and PEO. Before curing, the BA-a/PEO blends displayed the single and composition-dependant glass transition temperatures (Tg's) in the entire blend composition, and the equilibrium melting point depression was also observed in the blends. It is judged that the BA-a/PEO blends are completely miscible. The miscibility was mainly ascribed to the contribution of entropy to mixing free energy since the molecular weight of BA-a is rather low. However, phase separation occurred after curing reaction at the elevated temperature, which was confirmed by differential scanning calorimetry (DSC) and scanning electronic microscopy (SEM). It was expected that the PBA-a/PEO blends would be miscible since PBA-a possesses a great number of phenolic hydroxyls in the molecular backbone, which are potential to form the intermolecular hydrogen bonding interactions with oxygen atoms of PEO and thus would fulfill the miscibility of the blends. To interpret the experimental results, we investigated the variable temperature Fourier transform infrared spectroscopy (FTIR) of the blends via model compound. The FTIR results indicate that the phenolic hydroxyl groups could not form the efficient intermolecular hydrogen bonding interactions at the elevated temperatures (e.g. the curing temperatures), i.e. the phenolic hydroxyl groups existed mainly in the non-associated form in the system. Therefore, the decrease of the mixing entropy still dominates the phase behavior of thermosetting blends at the elevated temperature.

Functional alteration by NMDA antagonist: effects of L-Dopa, neuroleptics drug and postnatal administration.

Antiakinsic effects of the uncompetitive NMDA antagonists, memantine, amantadine and MK-801, and competitive antagonists, CGP 40116, alone or in co-administration with acute subthreshold dose of L-Dopa (5 mg/kg) in MPTP-treated mice, functional alterations induced by acute MK-801 in combinations with neuroleptic compounds or behavioural deficits following postnatal administration of MK-801 were investigated. Memantine and amantadine injected 60 min before the subthreshold dose of L-Dopa (5 mg/kg), induced antiakinesic actions in hypokinesic MPTP-treated mice. Concurrently, higher doses of memantine and MK-801 caused dyskinesic changes, reducing further rearing (10 and 30 mg/kg) and locomotor (30 mg/kg) behaviour of the MPTP mice; MK-801 elevated locomotion (0.1 mg/kg) but reduced rearing (0.3 mg/kg). In control, saline-treated mice, memantine (3, 10 and 30 mg/kg) and MK-801 (0.1 and 0.3 mg/kg) increased locomotor behaviour but decreased rearing behaviour. In rats, MK-801 induced marked increases in locomotor activity and disruptions of circular swim maze acquisition that were to greater or lesser extents blocked or potentiated by neuroleptic compounds: SCH 23390 (0.005 and 0.05 mg/kg) and clozapine (5.0 and 10.0 mg/kg) dose-dependently antagonised MK-801 (0.3 mg/kg) induced locomotor activity whereas raclopride (0.1 mg/kg) and haloperidol (0.1 mg/kg) attenuated it dose-specifically. Amperozide (0.5 mg/kg) attenuated the MK-801 effect but potentiated it at the 2.0 mg/kg dose. In the circular swim maze, raclopride (0.01 mg/kg) and SCH 23390 (0.05 mg/kg) improved the acquisitive performance of rats administered MK-801 (0.03 mg/kg) acutely whereas clozapine (10.0 mg/kg) and amperozide (2.0 mg/kg) deteriorated the performance of MK-801-treated rats. Postnatal administration of MK-801 (0.05 mg/kg, day 11 after birth) induced severe functional alterations in adult mice. At 70 days of age, MK-801 mice showed an initial hypoactivity followed by marked hyperactivity in the motor activity test chambers. These mice showed deficits in habituation, a nonassociative form of learning. Their hyperactivity in the test chambers was reversed by a low dose of d-amphetamine (0.25 mg/kg). Taken together, these findings display a wide range of acute/long-term functional alterations induced by NMDA antagonists, particularly MK-801, associated with animal models of brain disorders.

Noncovalent association with stress protein facilitates cross-priming of CD8+ T cells to tumor cell antigens by dendritic cells.

A viral oncogene carrying well-defined K(b)/D(b)-restricted epitopes was expressed in a heat shock protein (hsp)-associated or nonassociated form in the murine tumor cells P815 and Meth-A. Wild-type SV40 large T-Ag (wtT-Ag) is expressed without stable hsp association; mutant (cytoplasmic cT-Ag) or chimeric (cT272-green fluorescent fusion protein) T-Ag is expressed in stable association with the constitutively expressed, cytosolic hsp73 (hsc70) protein. In vitro, remnants from apoptotic wtT-Ag- or cT-Ag-expressing tumor cells are taken up and processed by immature dendritic cells (DC), and the K(b)/D(b)-binding epitopes T1, T2/3, and T4 of the T-Ag are cross-presented to CTL in a TAP-independent way. DC pulsed with remnants of transfected, apoptotic tumor cells cross-presented the three T-Ag epitopes more efficiently when they processed ATP-sensitive hsp73/cT-Ag complexes than when they processed hsp-nonassociated (native) T-Ag. In vivo, more IFN-gamma-producing CD8+ T cells were elicited by a DNA vaccine that encoded hsp73-binding mutant T-Ag than by a DNA vaccine that encoded native, non-hsp-binding T-Ag. Three- to 5-fold higher numbers of T-Ag (T1-, T2/3-, or T4-) specific, D(b)/K(b)-restricted IFN-gamma-producing CD8+ T cells were primed during the growth of transfected H-2(d) Meth-A/cT tumors than during the growth of transfected Meth-A/T tumors in F(1)(b x d) hosts. Hence, the association of an oncogene with constitutively expressed, cytosolic hsp73 facilitates cross-priming in vitro and in vivo of CTL by DC that process material from apoptotic cells.

Functional deficits following neonatal dopamine depletion and isolation housing: circular water maze acquisition under pre-exposure conditions and motor activity.

Seven experiments and several behavioural tests were performed to study the effects of housing condition and experimental test conditions upon the behavioural responses and performance of adult rats neonatally treated with 6-hydroxydopamine (6-OHDA, 100 ug intracisternally, i.c.) or with vehicle. Postnatal 6-OHDA induced locomotor and total activity hyperactivity and deficits in navigational learning in a circular swim maze that were blocked by pretreatment with a dopamine (DA) reuptake inhibitor but not a noradrenaline (NA) reuptake inhibitor. Isolation-housing induced deficits in maze learning performance. Grouped housing improved the maze learning performance of 6-OHDA treated rats whereas vehicle treated rats that were isolation housed performed better following latent learning (LL) pre-exposure trials. 6-OHDA treated rats that received both Grouped housing and latent learning trials performed better on the spatial navigation task than those that received Grouped housing but no latent learning or Isolation housing and latent learning. Analysis of habituation quotients indicated marked deficits by 6-OHDA-treated rats suggesting inability to acquire this simple, nonassociative form of learning. Methylphenidate increased all three parameters of motor activity: locomotion, rearing and total activity, in both Isolation-housed and Group-housed rats from 60- to 90- or 120-min post-injection. NDO 008 induced variable and parameter-dependent effects: locomotion was elevated initially in both Isolated and Grouped rats by the compound and then reduced in the Isolated rats only whereas total activity was only elevated initially in the Isolated rats and unaffected in the Grouped rats. Rearing behaviour was reduced markedly, directly post-injection, in the Isolation-housed rats. DA, DOPAC and HVA concentrations in the striatum, nucleus accumbens, olfactory tubercle and midbrain were reduced but most markedly in the striatum. 5-Hydroxytryptamine (5-HT) and 5-hydroxyindole acetic acid (5-HIAA) concentrations were elevated in the striatum, nucleus accumbens (not 5-HIAA) and olfactory tubercle.

Book Reviews

Book Reviews

The concepts of 'sameness' and 'difference' in an insect.

Insects process and learn information flexibly to adapt to their environment. The honeybee Apis mellifera constitutes a traditional model for studying learning and memory at behavioural, cellular and molecular levels. Earlier studies focused on elementary associative and non-associative forms of learning determined by either olfactory conditioning of the proboscis extension reflex or the learning of visual stimuli in an operant context. However, research has indicated that bees are capable of cognitive performances that were thought to occur only in some vertebrate species. For example, honeybees can interpolate visual information, exhibit associative recall, categorize visual information and learn contextual information. Here we show that honeybees can form 'sameness' and 'difference' concepts. They learn to solve 'delayed matching-to-sample' tasks, in which they are required to respond to a matching stimulus, and 'delayed non-matching-to-sample' tasks, in which they are required to respond to a different stimulus; they can also transfer the learned rules to new stimuli of the same or a different sensory modality. Thus, not only can bees learn specific objects and their physical parameters, but they can also master abstract inter-relationships, such as sameness and difference.

Analyses of habituation in Caenorhabditis elegans.

Although the nonassociative form of learning, habituation, is often described as the simplest form of learning, remarkably little is known about the cellular processes underlying its behavioral expression. Here, we review research on habituation in the nematode Caenorhabditis elegans that addresses habituation at behavioral, neural circuit, and genetic levels. This work highlights the need to understand the dynamics of a behavior before attempting to determine its underlying mechanism. In many cases knowing the characteristics of a behavior can direct or guide a search for underlying cellular mechanisms. We have highlighted the importance of interstimulus interval (ISI) in both short- and long-term habituation and suggested that different cellular mechanisms might underlie habituation at different ISIs. Like other organisms, C. elegans shows both accumulation of habituation with repeated training blocks and long-term retention of spaced or distributed training, but not for massed training. Exposure to heat shock during the interblock intervals eliminates the long-term memory for habituation but not the accumulation of short-term habituation over blocks of training. Analyses using laser ablation of identified neurons, and of identified mutants have shown that there are multiple sites of plasticity for the response and that glutamate plays a role in long-term retention of habituation training.

Context conditioning in habituation in the nematode Caenorhabditis elegans.

Habituation has traditionally been considered a nonassociative form of learning. However, recent research suggests that retention of this nonassociative form of learning may be aided by associations formed during training. An example of this is context conditioning, in which animals that are trained and tested in the presence of a contextual cue show greater retention than animals trained and tested in different environments. This article reports context conditioning in habituation in the nematode Caenorhabditis elegans. The results showed that retention of habituation to tap at both 10- and 60-s interstimulus intervals was significantly greater if training and testing occurred in the presence of the same chemosensory cue (NaCH3COO). This context conditioning showed both extinction and latent inhibition, demonstrating that these simple worms with only 302 neurons are capable of associative context conditioning.