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PubMed2007

Demographic review of a captive colony of callitrichids (Callithrix kuhlii).

Although reports on colony demographics for a variety of callitrichid species are available in the literature, to date there has not been a detailed examination of Wied's black tufted-ear marmoset (Callithrix kuhlii). The purpose of this study is to present colony demographics for C. kuhlii from the University of Nebraska at Omaha's Callitrichid Research Center from 1991 to 2002. C. kuhlii are currently held in a number of zoological parks in the United States and abroad; however, the University of Nebraska at Omaha held the only breeding colony in North America. Here we report data on lifespan, sex ratio, litter size, and interbirth interval (IBI) for that captive breeding colony.

Animals↗

PubMed1993

Brief communication: gut morphology of Callithrix nigriceps and Saguinus labiatus from western Brazilian Amazonia.

Quantitative data on the gut proportions of the recently-described black-headed marmoset (Callithrix nigriceps) from western Brazilian Amazonian are presented. Like other marmosets, C. nigriceps exhibits a relatively large caecum and colon associated with adaptations for gummivory. The degree of gut specialisation recorded is greater than that for any other platyrrhine for which quantitative data are available. Gum was found in the stomach contents of all four individuals examined, although the fruit/seeds of as many as seven different plant taxa were also encountered, in addition to insects of six different orders. Gut proportions of two Saguinus labiatus from the same region were typical of the more frugivorous tamarins, and stomach contents included only fruit and insects. Relative gut proportions of both species were closely similar to those of congeners from the same region.

Animals↗

PubMed2002

Cortical and thalamic connections of the parietal ventral somatosensory area in marmoset monkeys (Callithrix jacchus).

Microelectrode mapping methods were used to define the parietal ventral somatosensory area (PV) on the upper bank of the lateral sulcus in five marmosets (Callithrix jacchus). In the same animals, neuroanatomical tracers were placed into electrophysiologically identified sites in PV and/or the second somatosensory area (S2). Foci of anterograde and retrograde label were related to electrophysiological maps of cortical areas and cortical and thalamic architecture. The results lead to the following conclusions: (1) Multiunit recordings from cortex on the upper bank of the lateral sulcus demonstrate that PV is somatotopically organized, with the face representation adjoining area 3b and the hindlimb and tail representations away from this border in cortex deep on the upper bank of the lateral sulcus. The forelimb representation is caudal in PV adjacent to the S2 forelimb representation. The body surface representation in PV approximates a mirror image of that in S2; (2) Areas PV and S2 are less myelinated and have less cytochrome oxidase enzyme activity than area 3b; (3) The ventroposterior inferior nucleus (VPI) of the thalamus provides the major somatosensory projections to PV. PV is reciprocally connected with VPI and anterior pulvinar; (4) PV has ipsilateral cortical connections with areas 3a, 3b, 1, and M1 and higher order somatosensory fields, and at least most of these connections are somatotopically matched; and (5) Callosal connections of PV are with S2 and PV of the other cerebral hemisphere. These results further establish PV as one of at least four somatosensory areas of the lateral sulcus of primates.

Animals↗

PubMed2005

Resolving the organization of the New World monkey third visual complex: the dorsal extrastriate cortex of the marmoset (Callithrix jacchus).

We tested current hypotheses on the functional organization of the third visual complex, a particularly controversial region of the primate extrastriate cortex. In anatomical experiments, injections of retrograde tracers were placed in the dorsal cortex immediately rostral to the second visual area (V2) of New World monkeys (Callithrix jacchus), revealing the topography of interconnections between the "third tier" cortex and the primary visual area (V1). The data indicate the presence of a dorsomedial area (DM), which represents the entire upper and lower quadrants of the visual field, and which receives strong, topographically organized projections from the superficial layers of V1. The visuotopic organization and boundaries of DM were confirmed by electrophysiological recordings in the same animals and by architectural characteristics which were distinct from those found in ventral extrastriate cortex rostral to V2. There was no electrophysiological or histological evidence for a transitional area between V2 and DM. In particular, the central representation of the upper quadrant in DM was directly adjacent to the representation of the horizontal meridian that marks the rostral border of V2. The present results argue in favor of the hypothesis that the third visual complex in New World monkeys contains different areas in its dorsal and ventral components: area DM, near the dorsal midline, and a homolog of area 19 of other mammals, located more lateral and ventrally. The characteristics of DM suggest that it may correspond to visual area 6 (V6) of Old World monkeys.

Action Potentials↗

PubMed2005

Functional organization and hemispheric comparison of primary auditory cortex in the common marmoset (Callithrix jacchus).

Hemispheric fine-grain maps of primary auditory cortex (AI) were derived from microelectrode penetrations in the temporal gyrus of the common marmoset (Callithrix jacchus) to 1) compare the functional organization of AI in the marmoset with other mammalian species and 2) compare the right and left AI maps in individual monkeys. Frequency receptive fields (FRFs) were recorded with pure tones. Five FRF parameters were analyzed: characteristic frequency, threshold, sharpness of tuning 10 dB and 40 dB above threshold, and minimum response latency. The present study confirms that the functional organization of AI is characterized by a robust tonotopic frequency gradient overlaid with spatially clustered distributions of other FRF parameters. Globally, this functional organization of AI in the common marmoset is similar to that in other mammalian species. With respect to within-subject hemispheric comparisons of the five FRF parameters, a coherent pattern of asymmetry is not evident in marmoset AI. The overall results indicate that the basic functional organization between hemispheres is similar but not identical.

Acoustic Stimulation↗

PubMed2006

Development of the neural retina and its vasculature in the marmoset Callithrix jacchus.

The morphological sequence of retinal development in the New World marmoset monkey Callithrix jacchus is similar to previous reports in Macaca and humans. The incipient fovea is present at fetal day (Fd) 100 as the only part of the retina that contains five distinct layers, including a single layer of cone photoreceptors. A foveal pit begins to form at Fd 135 in the center of the foveal avascular zone which is surrounded by a ring of blood vessels (BV) and astrocytes. At birth (Fd 144) the fovea has a single layer of cones over the pit center where the inner retinal layers are thinned but still separated. After birth the fovea rapidly matures so that foveal cone and pit morphology are similar to adult by 4 months. Five distinct layers and the BV plexus in the nerve fiber layer are present to the retinal edge in neonatal marmosets. Near the optic disc BV are sprouting into outer retinal layers at birth and vascularization of the outer retina is completed by 2 to 3 months. Retinal length increases sharply up to Fd 135, but undergoes a quiescent period around birth during which pit formation begins. Length then increases again up to 4mo, followed by a slow increase into adulthood. The postnatal increase is accompanied by a marked thinning of the peripheral retina. The pars plana appears after birth and its length increases at least until 2 years of age. The major difference between marmoset and Macaca is the relative immaturity of the marmoset fovea at birth, and its rapid development after birth. This makes the marmoset a good candidate for neonatal experimental manipulation of retinal and eye development.

Animals↗

PubMed1985

A callosal projection of area 17 upon the border region of area MT in the marmoset monkey, Callithrix jacchus.

Injections of the retrograde tracer HRP into the border region of the temporal visual area MT and adjoining cortex in Callithrix labeled pyramidal neurons in area 17 of the contralateral hemisphere. Evidence is presented that this newly discovered heterotopic callosal projection of the monkey striate cortex connects regions of representation of the zero vertical meridian of the visual field in a retinotopic order.

Animals↗

PubMed1986

Frequency representation in auditory cortex of the common marmoset (Callithrix jacchus jacchus).

The location and characteristics of the primary auditory cortex of the common marmoset, Callithrix jacchus jacchus, were determined in five anesthetized male adult animals by mapping the responses of cortical units and unit clusters to pure tone stimuli presented to the contralateral ear. The primary auditory cortex lies largely ventral to the lateral sulcus, the only major fissure on the lateral cortex of this smooth-brained primate, but in some animals it may extend significantly down the ventral bank of this sulcus. Responses are distributed such that low best frequencies are found rostroventrally whereas high best frequencies occur caudally. The disposition of frequency-band contours is fan-shaped, with contours separating low-frequency octaves nearly parallel to the lateral sulcus and high-frequency (greater than 8 kHz) contours perpendicular to that sulcus. Best frequencies range from 0.6 to 30 kHz across the primary field, but there is a disproportionate representation of the three octaves between 2 and 16 kHz. The most sensitive thresholds (as low as -2 dB SPL) are found between 7 and 9 kHz. The primary auditory cortex is similar in cytoarchitecture to that reported for the cat, showing a blurring of lamination in the middle layers (II-IV) and a preponderance of small cells in these merged layers, giving a highly granular appearance. The accessibility of the cochlear representation on the gyral surface makes the marmoset an attractive animal for studies of primate auditory cortex.

Animals↗

PubMed1988

Postnatal development of dendrites of relay neurons in the lateral geniculate nucleus of the marmoset (Callithrix jacchus): a quantitative Golgi study.

Dendrites of multipolar relay neurons in the lateral geniculate nucleus of the marmoset (Callithrix jacchus), at various ages from birth to adulthood, were studied in rapid Golgi preparations. The dendrites were analyzed by means of three-dimensional computer reconstructions and decomposed into intermediate and terminal segments, both of which were further classified according to their centrifugal order. Measurements were made of the number of segments per dendrite, the total length of dendrites, and the mean length of intermediate and terminal segments. In adult marmosets, there are four stem dendrites on average per neuron, and each dendrite divides into a mean of 14 segments. Between birth and 6 weeks of age, the mean dendritic length doubles, mainly because of changes in terminal segments. There is a significant decrease in dendritic length into adulthood. The total number of stem dendrites does not change after birth, but during the first postnatal week dendrites lose distal segments, after which there is a significant increase in the number of segments of orders 3 to 7. The mean length of intermediate segments does not change with age, nor with order, whereas the length of terminal segments increases from 50 to 120 microns from birth to 6 weeks of age, and then decreases to the adult value of 80 microns. In conclusion, during the period of most rapid visual development, important morphological changes occur in geniculate relay-cell dendrites, involving essentially terminal segments. These observations correlate well with changes of geniculate volume and neuronal density.

Animals↗

PubMed1994

Distribution of cytochrome oxidase and parvalbumin in the primary visual cortex of the adult and neonate monkey, Callithrix jacchus.

The anatomical distributions of the mitochondrial enzyme cytochrome oxidase (CO) and of the calcium binding protein parvalbumin (PV) were studied in the striate cortex of adult and neonate New World monkeys (Callithrix jacchus). In the adult marmoset, both proteins were found in laminar arrangements similar to those described for the macaque monkey, with prominent bands of PV-like immunoreactive (PV-LI) puncta in layers IV and IIIb, and fairly evenly distributed PV-LI nonpyramidal neurons. Furthermore, the pattern of CO activity in area 17 of the neonate marmoset was almost identical to the CO pattern described in neonate macaque and squirrel monkeys. It came, therefore, as a surprise to find that the adult pattern of PV-like immunoreactivity (PV-LI) in the marmoset striate cortex arises from a neonatal pattern strikingly different from that seen in any developmental stage of the macaque, or in any other mammal studied so far. In the deep layers IV through VI of the neonate marmoset, a large number of PV-LI neurons was stained in bandlike patterns, their number in layers IV and V exceeding the number of PV-LI neurons present in these layers of the adult marmoset area 17. Staining of layers IV and VI was restricted to area 17 and involved nonpyramidal cells and their processes. The stained band of layer V, in contrast, continued throughout most of the neocortex. In area 17, an estimated 10 to 20% of the stained cells in layer V exhibited pyramidal shapes. The findings show that the expression of PV by visual cortical cells occurs before birth and suggest that the comparatively early onset of PV expression is not dependent on the onset of textured vision. The exuberant number of stained cells in some layers, and particularly the staining of pyramidal cells, in the neonate marmoset, suggest that a considerable number of cells possesses the stainability for PV-LI only transiently, i.e., in the marmoset, these cells have a specific demand for parvalbumin during this phase of their development.

Animals↗

PubMed2005

Micronucleated erythrocyte frequencies in old and new world primates: measurement of micronucleated erythrocyte frequencies in peripheral blood of Callithrix jacchus as a model for evaluating genotoxicity in primates.

Nonhuman primates are of particular relevance in evaluating the potential toxicity of drugs and environmental agents. We have used previously published information and data from the present study to establish a relationship for New World (NW) and Old World (OW) primates on the basis of the frequency of spontaneous micronucleated erythrocytes (MNEs) observed in peripheral blood. Data on spontaneous MNEs in peripheral blood from 15 species of primates, including humans, indicate that NW primates have significantly (P < 0.01) higher MNE frequencies (group mean, 9.5 +/- 7.3 MNEs/10,000 erythrocytes; range, 0.7-20.5/10,000 erythrocytes) than OW primates (group mean, 1.0 +/- 0.9 MNEs/10,000 erythrocytes; range, 0.0-2.6 MNEs/10,000 erythrocytes). Humans are believed to have developed in the OW, and human MNE frequencies were similar to those described for OW primate species. We selected the common marmoset (Callithrix jacchus), a NW primate, to determine whether therapeutic pediatric doses of Metotrexate (MTX; 2.5 mg/kg), Cyclophosphamide (CP; 5 mg/kg), Cytosine-arabinoside (Ara-C; 3 mg/kg), or 5-Fluorouracil (5-FU; 10 mg/kg), administered daily for two consecutive days, increase the frequency of micronuclei. Micronucleated polychromatic erythrocyte frequencies were increased significantly in groups receiving MTX, CP and Ara-C, while MNE frequencies were increased by the Ara-C treatment. The results of this study indicate that NW primates have higher spontaneous MNE frequencies than OW primates, and because of this, NW primates like the common marmoset, may be suitable for evaluating the genotoxicity of chemical agents.

Animals↗

PubMedDate not supplied

Preclinical safety studies with recombinant human interleukin 6 (rhIL-6) in the primate Callithrix jacchus (marmoset): comparison with studies in rats.

The haemopoietic and immunostimulatory properties of recombinant human interleukin 6 (rhIL-6) might be used clinically in cancer patients. For the preclinical assessment of the safety of such a therapy, we chose the primate marmoset (Callithrix jacchus) and Wistar rats. Recombinant hIL-6 given to marmosets at doses of up to 1000 micrograms kg-1 day-1 over 4 and 9 weeks did not induce fever and was very well tolerated. Haematological alterations included a sustained two- to threefold increase of thrombocyte counts, peaking at 4 weeks, as well as an increase in neutrophils and basophils. The number of bone marrow megakaryocytes at 4 and 9 weeks was not increased, but the ploidy grade was augmented. An acute-phase protein response was observed within 24 h after the first IL-6 administration, which reached a maximum after 1 week. The acute-phase protein response was not accompanied by functional or morphological signs of hepatocellular damage. Increased immunoglobulin and soluble IL-2 receptor in the serum levels reflected systemic immunostimulation. Recombinant hIL-6 was also given to rats at 500 micrograms kg-1 day-1 s.c. for 4 weeks, where it induced a stimulation of thrombopoiesis associated with increased platelet counts within 1 week. Furthermore, rhIL-6-treated rats had signs of immunostimulation and increased acute-phase reactants in serum, as in marmosets. There was no evidence of renal glomerular or hepatic pathology.(ABSTRACT TRUNCATED AT 250 WORDS)

Acute-Phase Proteins↗

PubMed1985

Attenuated Junin virus infection in Callithrix jacchus.

Twenty marmosets, male Callithrix jacchus, were used during this study. Fifteen of the marmosets were inoculated with 5,000 TCID50 of the attenuated XJC13 strain of Junin virus by intramuscular route and five were left as uninoculated controls. Animals were observed for a 420-day period. In order to carry out virologic, hematologic, serologic, and histologic studies the animals were bled and/or killed at different days post infection(pi). Results obtained showed that the attenuated strain produced an infection with no mortality or signs of illness. There was only a slight loss of weight at 18-40 days pi, which was soon recovered. Viremia was present from day 6 to 22, titers peaking at 4.0 log. Viral spread was limited to the lungs, spleen, lymph nodes, and bone marrow in the animal killed on day 14. No virus was found in the organs of the animal killed on day 23, and neither hematologic alterations nor pathologic lesions were seen in these monkeys except for ganglionar hypertrophy with immunoblast proliferation. Antigen was detected by immunofluorescence (IF) in lymph nodes, spleen, adrenals, lungs and brain. Neutralizing antibodies were detected from the third week onward. Protection conferred by the XJC13 strain proved effective when XJC13-inoculated monkeys were challenged with 1,000 TCID50 of the pathogenic XJ strain at days 60 or 380 pi, while normal controls died. When viral persistence was searched for on days 370, 390, and 420 pi, no infectious virus was detected, but viral antigen was seen in certain organs, which, however, lacked tissue damage.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

PubMed1995

Chronic L-DOPA administration induces dyskinesias in the 1-methyl-4- phenyl-1,2,3,6-tetrahydropyridine-treated common marmoset (Callithrix Jacchus).

Dyskinesias occur in the majority of patients with Parkinson's disease chronically treated with L-DOPA and also occur in several nonhuman primate species after 1-methyl-4phenyl-1,2,3,6-tetrahydropyridine (MPTP) and L-DOPA treatment. The common marmoset (Callithrix jacchus) shows parkinsonian motor deficits after MPTP administration, and we now report dyskinesias occurring in this species during chronic L-DOPA exposure. Marmosets rendered chronically parkinsonian after MPTP administration were treated orally with L-DOPA plus carbidopa for 3 weeks. After several days the animals began to display chorea, choreoathetosis, and dystonia. The severity of dyskinesias varied between the animals, with the most severely parkinsonian animals displaying the most dyskinetic movements. Each animal showed an idiosyncratic pattern of dyskinesias, which was highly reproducible. These L-DOPA-primed animals also received other D2 D1, and mixed D1/D2 agonist drugs. Quinpirole, bromocriptine, pergolide, apomorphine, and A-77636 all produce dyskinesias that were identical in character to those seen after L-DOPA administration, but the D1 agonist A-77636 gradually abolished dyskinesias while preserving its antiparkinsonian activity. The MPTP-treated marmoset provides a useful model in which to study dyskinesias in Parkinson's disease and to examine new therapeutic strategies aimed at alleviating this common side effect of chronic dopamine replacement therapy.

1-Methyl-4-phenyl-1,2,3,6-tetrahydropyridine↗

PubMed2005

Ejaculates from the common marmoset (Callithrix jacchus) contain semenogelin and beta-microseminoprotein but not prostate-specific antigen.

Human seminal plasma contains high concentrations of prostatic acid phosphatase (PAP), prostate-specific antigen (PSA), beta-microseminoprotein (MSP), semenogelin I (SgI), and semenogelin II (SgII), whereas only PAP and MSP are present in rodents. In order to gain a better understanding of the evolution and function of semen proteins, we have studied ejaculates from the common marmoset (Callithrix jacchus)-a New World monkey. Semen samples were analyzed with SDS-PAGE, Western blotting, and isoelectric focusing. Under reducing conditions the dominating protein components appear as heterogeneous material of 55-70 kDa and distinct protein bands of 85, 17, 16, and 15 kDa. The heterogeneous material contains glycosylated material detected by an antiserum recognizing both human SgI and SgII. Southern blotting indicates that the common marmoset has genes for both SgI and SgII. There are several marmoset MSP genes, but the strong immunoreactivity against one 15 kDa semen component with pI 7.3 suggests preferential expression of one gene in the prostate. Expression of two other genes cannot be excluded as indicated by weak reaction to isoforms with pI 6.6 and 4.9. Unexpectedly, PSA was not detected by either immunological methods or activity measurements. This is in agreement with results from Southern blotting suggesting that the common marmoset might not have a PSA gene. Thus, in this study we have shown that semen coagulum proteins are present in marmoset seminal plasma, but the lack of PSA precludes a similar liquefaction as of human semen.

Animals↗

PubMed1994

Embryotoxic effects of thalidomide derivatives in the non-human primate callithrix jacchus. IV. Teratogenicity of micrograms/kg doses of the EM12 enantiomers.

The dose-response of the teratogenic potency of the thalidomide (Thd) derivative EM12 was evaluated in the common marmoset (Callithrix jacchus). The smallest daily dose found to be effective was 30 micrograms EM12/kg body wt. This is the lowest dose of a Thd derivative ever reported to induce severe skeletal abnormalities. Ten micrograms EM12/kg body wt may be considered the no-observed-adverse-effect-level (NOAEL) under the experimental conditions chosen. The teratogenic potencies of the two EM12 enantiomers were tested at 100 micrograms/kg body wt, the dose which just induces an almost 100% effect in the case of the racemate. The S(-)-EM12 was found to induce typical severe limb abnormalities such as amelia, phocomelia, and radius aplasia, and none of the exposed fetuses were devoid of skeletal defects. In contrast, only few and minor skeletal defects were observed after application of the R(+) enantiomer. Although a pronounced teratogenic potency of the R(+)-EM12 can now largely be excluded, these low-dose studies are not sufficient to completely rule out any teratogenic potential of this enantiomer, since racemisation to small amounts of the S(-) form may occur in vivo. Further studies with Thd derivatives which are unable to racemise are necessary to prove the assumed complete ineffectiveness of the R(+) enantiomers.

Abnormalities, Drug-Induced↗

PubMed2000

Cyto- and chemoarchitecture of basal forebrain cholinergic neurons in the common marmoset (Callithrix jacchus).

The cyto- and chemoarchitecture of basal forebrain cholinergic neurons (BFCN) was investigated in the lower primate, the common marmoset (Callithrix jacchus). A large population of magnocellular, hyperchromic, and choline acetyltransferase (ChAT)-positive neurons was detected in the marmoset basal forebrain. The distribution of these neurons was similar to those in higher primates. Thus, ChAT-positive neurons were observed in the medial septum (Ch2), the vertical (Ch2) and horizontal (Ch3) limbs of the diagonal band of Broca, and the nucleus basalis of Meynert (Ch4). The Ch4 complex was relatively well differentiated and displayed distinct sectors. We detected anterior (Ch4a, with a medial and a lateral subdivision), intermediate (Ch4i, with a dorsal and a ventral subdivision), and posterior (Ch4p) sectors in the marmoset Ch4. The Ch4i was relatively small while the Ch4p was large. Similar to the rodent, the marmoset Ch1 extended quite a distance posteriorly, and the Ch4p displayed a major interstitial component distributed within the globus pallidus, its medullary laminae, and the internal capsule. Virtually all of the marmoset BFCN displayed acetylcholinesterase activity, and low affinity (p75(NTR)) and high affinity (Trk) neurotrophin receptor immunoreactivity. A majority contained immunoreactivity for calbindin-D(28K) and calretinin. Many of the Ch4 neurons also displayed tyrosine hydroxylase immunoreactivity. The BFCN lacked galanin immunoreactivity, but were innervated by galanin-positive fibers. None of the marmoset BFCN were NADPH-d-positive. Thus, the BFCN display major anatomical and biochemical differences in the marmoset when compared with higher primates. The marmoset BFCN also display many characteristics common to other primates. This fact, combined with the relatively short life span of the marmoset, indicates that this species may be ideal for studies of age-related changes in the BFCN.

Acetylcholinesterase↗

PubMed2003

Both short- and long-acting D-1/D-2 dopamine agonists induce less dyskinesia than L-DOPA in the MPTP-lesioned common marmoset (Callithrix jacchus).

The current concept of dyskinesia is that pulsatile stimulation of D-1 or D-2 receptors by L-DOPA or short-acting dopamine agonists is more likely to induce dyskinesia compared to long-acting drugs producing more continuous receptor stimulation. We now investigate the ability of two mixed D-1/D-2 agonists, namely pergolide (long-acting) and apomorphine (short-acting), to induce dyskinesia in drug-nai;ve MPTP-lesioned primates, compared to L-DOPA. Adult common marmosets (Callithrix jacchus) were lesioned with MPTP (2 mg/kg/day sc for 5 days) and subsequently treated with equieffective antiparkinsonian doses of L-DOPA, apomorphine, or pergolide for 28 days. L-DOPA, apomorphine, and pergolide reversed the MPTP-induced motor deficits to the same degree with no difference in peak response. L-DOPA and apomorphine had a rapid onset of action and short duration of effect producing a pulsatile motor response, while pergolide had a slow onset and long-lasting activity producing a continuous profile of motor stimulation. L-DOPA rapidly induced dyskinesia that increased markedly in severity and frequency over the course of the study, impairing normal motor activity by day 20. Dyskinesia in animals treated with pergolide or apomorphine increased steadily, reaching mild to moderate severity but remaining significantly less marked than that produced by L-DOPA. There was no difference in the intensity of dyskinesia produced by apomorphine and pergolide. These data suggest that factors other than duration of drug action may be important in the induction of dyskinesia but support the use of dopamine agonists in early Parkinson's disease, as a means of delaying L-DOPA therapy and reducing the risk of developing dyskinesia.

1-Methyl-4-phenyl-1,2,3,6-tetrahydropyridine↗