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. 2014 Jun 2;8:56. doi: 10.3389/fncir.2014.00056

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Copyright © 2014 Hasegawa, Mukai, Asashima, Hojo, Ikeda, Komatsuzaki, Ooishi and Kawato.

This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) or licensor are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.

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Changes in the density and morphology of spines by activin and drugs in hippocampal slices. Spines were analyzed along the secondary dendrites of pyramidal neurons in the stratum radiatum of CA1 neurons. (A) Representative images of confocal micrographs; spines along dendrite without drug-treatments (Cont) and spines along dendrite after activin treatment for 2 h (Act). Maximal intensity projection onto XY plane from z-series confocal micrographs (MAX-XY), image analyzed by Spiso-3D (S) and 3 dimensional model (Model) are shown together. Bar, 3 μm. (B) Effect of treatments by activin and glutamate receptor blockers on the total spine density in CA1 neurons. Vertical axis is the average number of spines per 1 μm. A 2-h treatment in ACSF without drugs (Control, total spine numbers = 552, 8 neurons), with 10 ng/mL activin A (Act), with 10 ng/mL activin and 100 ng/mL follistatin (Act + Fol), with 10 ng/mL activin and 20 μM CNQX (Act + CNQX, total spine numbers = 977, 11 neurons, P = 0.005), with 10 ng/mL activin and 50 μM MK-801 (Act + MK). Statistical significance is calculated against activin treated group and indicated by stars. ^*P < 0.05, ^**P < 0.01. (C) Histogram of spine head diameters after a 2 h treatment in ACSF without drugs (Control, closed black diamond), with 10 ng/mL activin (closed red square), and with 10 ng/mL activin A and 100 ng/mL follistatin (closed blue diamond), with 10 ng/mL activin and 20 μM CNQX (closed green triangle), with 10 ng/mL activin A and 50 μM MK-801 (closed purple triangle). (D) Density of three subtypes of spines. Abbreviations are same as in (A). Vertical axis is the number of spines per 1 μm of dendrite. From left to right, small-head spines (Small), middle-head spines (Middle), and large-head spines (Large) type. ACSF without drugs (open column), Act (orange column), Act + Fol (blue column), Act + CNQX (green column), Act + MK801 (purple column) are shown. Statistical significance is calculated against activin treated group in each spine subtypes and comparisons reached significance are indicated by stars. The significance yielded P < 0.05. ^*P < 0.05, ^**P < 0.01. In (B,D), results are reported as mean ± s.e.m. For each drug treatment, we investigated 3 rats, 7 slices, 14 neurons, 28 dendrites and 1400–2000 spines. For control, we used 5 rats, 8 slices, 16 neurons, 31 dendrites and approx. 1700 spines.