STDP and BDNF in CA1 spines (Solinas et al. 2019)

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Accession:244412
Storing memory traces in the brain is essential for learning and memory formation. Memory traces are created by joint electrical activity in neurons that are interconnected by synapses and allow transferring electrical activity from a sending (presynaptic) to a receiving (postsynaptic) neuron. During learning, neurons that are co-active can tune synapses to become more effective. This process is called synaptic plasticity or long-term potentiation (LTP). Timing-dependent LTP (t-LTP) is a physiologically relevant type of synaptic plasticity that results from repeated sequential firing of action potentials (APs) in pre- and postsynaptic neurons. T-LTP is observed during learning in vivo and is a cellular correlate of memory formation. T-LTP can be elicited by different rhythms of synaptic activity that recruit distinct synaptic growth processes underlying t-LTP. The protein brain-derived neurotrophic factor (BDNF) is released at synapses and mediates synaptic growth in response to specific rhythms of t-LTP stimulation, while other rhythms mediate BDNF-independent t-LTP. Here, we developed a realistic computational model that accounts for our previously published experimental results of BDNF-independent 1:1 t-LTP (pairing of 1 presynaptic with 1 postsynaptic AP) and BDNF-dependent 1:4 t-LTP (pairing of 1 presynaptic with 4 postsynaptic APs). The model explains the magnitude and time course of both t-LTP forms and allows predicting t-LTP properties that result from altered BDNF turnover. Since BDNF levels are decreased in demented patients, understanding the function of BDNF in memory processes is of utmost importance to counteract Alzheimer’s disease.
Reference:
1 . Solinas SMG, Edelmann E, Leßmann V, Migliore M (2019) A kinetic model for Brain-Derived Neurotrophic Factor mediated spike timing-dependent LTP. PLoS Comput Biol 15:e1006975 [PubMed]
Model Information (Click on a link to find other models with that property)
Model Type: Neuron or other electrically excitable cell; Synapse; Dendrite;
Brain Region(s)/Organism: Hippocampus;
Cell Type(s): Hippocampus CA1 pyramidal GLU cell;
Channel(s): I Na,t; I_KD; I K; I h; I A; I Calcium;
Gap Junctions:
Receptor(s): AMPA; NMDA;
Gene(s):
Transmitter(s): Glutamate;
Simulation Environment: NEURON;
Model Concept(s): Facilitation; Long-term Synaptic Plasticity; Short-term Synaptic Plasticity; STDP;
Implementer(s): Solinas, Sergio [solinas at unipv.it]; Migliore, Michele [Michele.Migliore at Yale.edu];
Search NeuronDB for information about:  Hippocampus CA1 pyramidal GLU cell; AMPA; NMDA; I Na,t; I A; I K; I h; I Calcium; I_KD; Glutamate;
# Records
spine_records = [
    {'variable':'cai','location':0.5,'unit':'mM'}
    ,{'variable':'gluti','location':0.5,'unit':'mM'}
    ,{'variable':'v','location':0.5,'unit':'mV'}
    ,{'variable':'ica','location':0.5,'unit':'mM'}
    ,{'variable':'iampa','point_process':'AMPA','location':0.5,'unit':'nA'}
    ,{'variable':'U_SE_factor','point_process':'AMPA','location':0.5,'unit':''}
    ,{'variable':'U_SE','point_process':'AMPA','location':0.5,'unit':''}
    ,{'variable':'inmda','point_process':'NMDA','location':0.5,'unit':'nA'}
    ,{'variable':'ica','point_process':'NMDA','location':0.5,'unit':'nA'}
    ,{'variable':'mgb','point_process':'NMDA','location':0.5,'unit':'nA'}
    ,{'variable':'iglut','location':0.5,'unit':'mM'}
    ,{'variable':'RM','point_process':'RMECB','location':0.5,'unit':'mM'}
    ,{'variable':'RMr','point_process':'RMECB','location':0.5,'unit':'mM'}#    ,{'variable':'RMBLKe','point_process':'RMECB','location':0.5,'unit':'??'}
    ,{'variable':'RMBLK','point_process':'RMECB','location':0.5,'unit':'mM'}
    ,{'variable':'post_intra','point_process':'RMECB','location':0.5,'unit':'mM'}
    ,{'variable':'proBDNF','point_process':'BDNF','location':0.5,'unit':'mM'}
    ,{'variable':'fused_vesicles','point_process':'BDNF','location':0.5,'unit':'#'}
    ,{'variable':'mBDNF','point_process':'BDNF','location':0.5,'unit':'mM'}
    ,{'variable':'PC','point_process':'BDNF','location':0.5,'unit':'mM'}
    ,{'variable':'TrkB','point_process':'BDNF','location':0.5,'unit':'mM'}
    ,{'variable':'intracell_signaling','point_process':'BDNF','location':0.5,'unit':'mM'}
    ,{'variable':'g_factor','point_process':'AMPA','location':0.5,'unit':''}
    ]

Vrest = -70 # mV
Rm = 26000
RmDend = Rm
RaAll= 150
CmDend = 1.4
gna = 0.07
nash = 0
gkdr = 0.055
kdrsh = 0
gc = 1.e-5
gcalbar = 0.5
gcatbar = 0.2
gcanbar = 0.5
temperature = 35 # deg
erev_na = 55 # mV
erev_k = -90 # mV
check = False #True

IC_delay_to_spike = 1.3 # ms
gc = 52e-5; # gives a BPAP [Ca]i amp = 1.71 uM, from 0.1 to 1.81 uM
nspines = 12
nspines_2 = 5
sp_delay_env = 8

spine_point_processes=[
    {'label':'RMECB',
     'type':'RM_eCB',
    'locations':[0.5],
    'parameters':{'tau_RM': 7e3, #1800e3, # next 10e3 #ms
                  'theta_cai_RM': 0.046, # mM this is tuned to distinguish between LTP1:4 deltat=5ms and LTP1:1 deltat=-5ms
                  'sigma_cai_RM' : 0.01e-3, # mM
                  'alpha_cai_RMBLK': 1.5e-2, # mM/ms
                  'theta_cai_RMBLK' : 0.12, # mM
                  'sigma_cai_RMBLK' : 0.0001, # mM
                  'tau_RMBLK' : 1e5, # mM
                  'tau_RMLTP11':1800e3*1.2, # ms
                  'alpha_RMru':1.1,#1.0,#0.54, 
                  'theta_RMru':0.15,
                  'sigma_RMru':0.001,
                  'tau_RM_RMr':1e3,
                  'alpha_RM_RMr':1,
                  'theta_RM_RMr': 0.02,
                  'sigma_RM_RMr': 0.001
                  }}, # to get 150% LTP as in Edlmann et al. 2015
    {'label':'AMPA',
     'type':'Wghkampa_preML',
    'locations':[0.5],
    'parameters':{'Pmax':4e-6,
                  'glut_factor': 40}},
    {'label':'BDNF',
     'type':'BDNF',
    'locations':[0.5],
    'parameters':{'theta_cai_BDNF': 0.1038,
                  'max_cai_BDNF': 0.16,
                  'max_BDNF_rel_delay' : 300e3,
                  'alpha_gAMPA': 1.5, #4.8, 
                  'theta_gAMPA': 0.01,#0.038,
                  'sigma_gAMPA': 0.00001,#0.0015,
                  'shift_gAMPA': 0,
                  'v_BDNF':0.002*1.0,
                  'proBDNF_fraction':0.3,
                  'duration_BDNF_release':1500e3}},
    {'label':'NMDA',
     'type':'ghknmda',
    'locations':[0.5],
    'parameters':{'Pmax':4e-6*4.5,
                  'mg':0.0001,
                  'mgb_k':0.22,
                  'Area': 1.0}}]
# Spike-EPSP envelop
spike_delay_rnd = 8

Block_RMBLK = 0

PPR = True
time_on_initialization = 100e3 # ms 600e3 = 10 min
time_to_begin_induction = 20e3 # ms
nstim = 70
induction_freq = 0.5 # Hz
BPAP_stimulus_amplitude = 5 # nA
BPAP_stimulus_amplitude_branch = 0.08 # nA
BPAP_dep_stimulus_duration = 2.5 # ms
BPAP_hyp_stimulus_duration = 0 # ms
time_after_induction = 0 # ms
test_freq = 0.05 # Hz = 1/(20sec)
time_start_induction = time_on_initialization + time_to_begin_induction
test_post_point_to_plot_interval = 2*60*1e3 # ms = 2 min see Fig1B of Edelmann2015
time_of_expression = 1800e3 # ms = 30 min
activate_LTP_protocol = True
# time_delta =  [-30,-20, -10, -5]
time_delta =  [ -5, -5, -5, -5]
# time_delta =  [-10, -10, -10, -10]
override_tstop = None

ca_tuning = False
if ca_tuning:
    time_on_initialization = 1e3 # ms
    time_to_begin_induction = 100 # ms
    nstim = 35
    induction_freq = 0.5 # Hz
    BPAP_stimulus_amplitude = 0.35 # nA
    BPAP_stimulus_amplitude_branch = 0.08 # nA
    BPAP_dep_stimulus_duration = 3 # ms
    BPAP_hyp_stimulus_duration = 0 # ms
    time_after_induction = 0 # ms
    test_freq = 0.05 # Hz = 1/(20sec)
    time_start_induction = time_on_initialization + time_to_begin_induction
    test_post_point_to_plot_interval = 2*60*1e3 # ms = 2 min see Fig1B of Edelmann2015
    time_of_expression = 1800e3 # ms = 30 min
    activate_LTP_protocol = True
    time_delta =  list(range(-15,16,5))#[-10, -5]#[-20, -15, -10, -5, 0, 5, 10, 15, 20] # ms [-10, -5, 10]#
    override_tstop = 5e3
    
data_file = 'store_multi'

# Settings for Figures
# Figure list
figure_size = (6,6)
panels = {'vm spine':(0.05,0,0.3,0.3),
          'cai':(0.05,0.4,0.3,0.3),
          'use':(0.7,0,0.3,0.3),
          'gfactor':(0.4,0.7,0.3,0.3),
          # 'Spine EPSP':(0.7,0,0.3,0.3),
          # 'Soma EPSP':(0.4,0.7,0.3,0.3),
          # 'fused vesicles':(0.7,0.4,0.3,0.3)
          }
panels_seq = ['vm spine','cai',
              #'Spine EPSP','Soma EPSP',
              'use','gfactor',#'use_trans',
              # 'fused vesicles',None,
              # 'rm','bdnf',
              ]

single_figures = {
    'vm spine':(0.05,0,0.3,0.3),
    'cai':(0.05,0.4,0.3,0.3),
    'Cont EPSP':(0.7,0,0.3,0.3),
    'Pot EPSP':(0.4,0.7,0.3,0.3),
    'U_SE':(0.7,0.4,0.3,0.3),#'fused vesicles':(0.7,0.4,0.3,0.3),
    'bdnf':(),
    'trkb':(),
    'use':(),
    'rm':(),
    'rmd':(),
    'gfactor':()
          }
summary_panels = {'EPSP_sl':[['#DC143C','d',2,14],['#00008B','s',1,14],['g','+',1,14]],
          'STDP':['#DC143C','#00008B']}
# panels = {'vm_spine':(4,3,1),
#           'cai':(4,3,4),
#           'Cont_EPSP':(4,3,7),
#           'Pot_EPSP':(4,3,8),
#           'fused_vesicles':(4,3,10)
#           }
paper_fig_panels = [
    'LTP','vm_branch_pre','vm_branch_post'
    ,'rm','bdnf'
    ]

one_spine = [
    # ['Spine_9/head_9','r'],
    # ['Spine_8/head_8','k'],
    # ['Spine_7/head_7','g'],
    # ['Spine_6/head_6','c'],
    # ['Spine_5/head_5','y'],
    # ['Spine_4/head_4','b'],
    # ['Spine_3/head_3','m'],
    # ['Spine_2/head_2','k'],
    # ['Spine_1/head_1','g'],
    ['Spine_0/head_0','k']
    ]
one_spine_soma = [
    # ['Spine_9/head_9','r'],
    # ['Spine_8/head_8','k'],
    # ['Spine_7/head_7','g'],
    # ['Spine_6/head_6','c'],
    # ['Spine_5/head_5','y'],
    # ['Spine_4/head_4','b'],
    # ['Spine_3/head_3','m'],
    # ['Spine_2/head_2','k'],
    # ['Spine_1/head_1','g'],
    ['Spine_0/head_0','k'],
    ['soma','k']]
all_spines = [
    ['Spine_17/head_17','lightgreen'],
    ['Spine_16/head_16','lightgreen'],
    ['Spine_15/head_15','lightgreen'],
    ['Spine_14/head_14','lightgreen'],
    ['Spine_13/head_13','darkgreen'],
    ['Spine_12/head_12','lightcoral'],
    ['Spine_11/head_11','coral'],
    ['Spine_10/head_10','lime'],
    ['Spine_9/head_9','lime'],
    ['Spine_8/head_8','salmon'],
    ['Spine_7/head_7','g'],
    ['Spine_6/head_6','c'],
    ['Spine_5/head_5','y'],
    ['Spine_4/head_4','b'],
    ['Spine_3/head_3','m'],
    ['Spine_2/head_2','gray'],
    ['Spine_1/head_1','k'],
    ['Spine_0/head_0','r'],
    ]
all_spines_soma = [
    ['Spine_9/head_9','lime'],
    ['Spine_8/head_8','salmon'],
    ['Spine_7/head_7','g'],
    ['Spine_6/head_6','c'],
    ['Spine_5/head_5','y'],
    ['Spine_4/head_4','b'],
    ['Spine_3/head_3','m'],
    ['Spine_2/head_2','gray'],
    ['Spine_1/head_1','k'],
    ['Spine_0/head_0','r'],

    ['soma','r']
    ]
    
figure_list = {
    'cai':{'title':'Spine $\mathrm{[Ca^{2+}]}$',
           'varnames':[['cai','k']],
           'hdf5_section':all_spines,
           'location':0.5,
           'legend':True,
           'time_window':[['first_induction',
                          -40,
                          +80]],
           'xunit':'ms',
           'xlabel':'Time',
           # 'ylim':[0,0.17],
           'ylabel':'Concentration',
           'thresholds':[spine_point_processes[0]['parameters']['theta_cai_RM'],
                         spine_point_processes[0]['parameters']['theta_cai_RMBLK'],
                         spine_point_processes[2]['parameters']['theta_cai_BDNF']]}
    ,'vm spine':{'title':'EPSP + BPAP',
           'varnames':[['v','k']],
           'hdf5_section':one_spine,
           'location':0.5,
           'xunit':'ms',
           'legend':False,
           'xlabel':'Time',
           'ylim':[-72,0],
           'ylabel':'Memb. pot.',
           'time_window':[['first_induction',
                          -40,
                          +80]]
                 }
    ,'Cont EPSP':{'title':'EPSP before LTP',
           'hdf5_section':all_spines_soma,
           'varnames':['vm'],
           'time_window':[['pre_induction',
                          -20,
                          +80]],
           'xlabel':'Time',
           'legend':False,
           'xunit':'ms',
           'ylabel':'Memb. pot.',
            'unit':'mV'}
    ,'Pot EPSP':{'title':'EPSP after LTP',
           'hdf5_section':all_spines_soma,
           'varnames':['vm'],
           'time_window':[['post_expression',
                          -20,
                          +80]],
           'xlabel':'Time',
           'xunit':'ms',
           'ylabel':'Memb. pot.',
            'unit':'mV'}
    ,'Spine EPSP':{'title':'EPSP at spine',
           'hdf5_section':one_spine,
           'varnames':['vm'],
           'time_window':[['pre_induction',
                          -20,
                          +80],
                          ['post_expression',
                          -20,
                          +80]],
           'xlabel':'Time',
           'legend':False,
           'xunit':'ms',
           'ylabel':'Memb. pot.',
            'unit':'mV'}
    ,'Soma EPSP':{'title':'EPSP at soma',
           'hdf5_section':[['soma','k']],
           'varnames':['vm'],
           'time_window':[['pre_induction',
                          -20,
                          +80],
                          ['post_expression',
                          -20,
                          +80]],
           'xlabel':'Time',
           'legend':False,
           'xunit':'ms',
           'ylabel':'Memb. pot.',
            'unit':'mV'}
    ,'fused vesicles':{'title':'Fused BDNF vesicles',
           'varnames':[['fused_vesicles','k']],
           'hdf5_section':all_spines,
           'location':0.5,
           'xunit':'min',
           'legend':False,
           'xlabel':'Time',
           'ylabel':'Count',
           # 'ylim':[0,70],
           'time_window':[['full',
                          -20,
                          +80]],
                       }
    ,'vm_branch_pre':{'title':'Memb. pot. in branch',
           'varnames':[['branch_base_v','Blues']],
           'hdf5_section':['pyr2005'],
           'location':0.5,
           'time_window':[[
               'pre_induction', # period
               2, # index
               -20, # pre time window
               +100]], # post time window
           'xunit':'ms',
           'legend':False,
           'xlabel':'Time',
           'ylabel':'Voltage'}
    # ,'vm_branch_post':{'title':'Memb. pot. in branch',
    #        'varnames':[['branch_base_v','Blues']],
    #        'hdf5_section':['pyr2005'],
    #        'location':0.5,
    #        'time_window':[[
    #            'post_expression', # period
    #            -10, # index
    #            -20, # pre time window
    #            +100]], # post time window
    #        'xunit':'ms',
           # 'legend':False,
    #        'xlabel':'Time',
    #        'ylabel':'Voltage'}
    ,'LTP':{'title':'Plasticity',
           'varnames':[],
           'hdf5_section':['pyr2005'],
           'xlabel':'Time',
           'legend':False,
           'ylabel':'Change',
            'unit':'%'}
    ,'STDP':{'title':'STDP',
           'varnames':[],
           'hdf5_section':['Spine_9/head_9'],
           'location':0.5,
           'time_window':[['first_induction',
                          -20,
                          +200]],
           'xunit':'ms',
           'legend':False,
           'xlabel':'Time',
           'ylabel':'Change',
            'unit':'%'}            
    ,'PPR':{'title':'PPR',
           'varnames':[],
           'hdf5_section':['Spine_9/head_9'],
           'location':0.5,
           'time_window':[['first_induction',
                          -20,
                          +200]],
           'xunit':'ms',
           'legend':False,
           'xlabel':'Time',
           'ylabel':'Change',
           'unit':'%'}            
    ,'rm':{'title':'Presynaptic plasticity chain',
           'varnames':[['RM','r'],['RMr','k']],
           'hdf5_section':all_spines,
           'xunit':'min',
           'location':0.5,
           'time_window':[['full',
                          -20,
                          +80]],
           'xlabel':'Time',
           'legend':False,
           'ylabel':'Concentration'}
    ,'fullrm':{'title':'Presynaptic plasticity chain',
           # 'varnames':[['RM','r'],['RMr','k'],['RMBLK','g'],['U_SE','b'],['post_intra','m']],
           'varnames':[['RM','r'],['RMr','k'],['post_intra','m']],
           'hdf5_section':all_spines,
           'xunit':'min',
           'location':0.5,
           'time_window':[['full',
                          -20,
                          +80]],
           'xlabel':'Time',
           'legend':False,
           'ylabel':'Concentration'}
    ,'rmd':{'title':'Presynaptic plasticity chain',
           'varnames':[['RMBLK','g']],
           'hdf5_section':all_spines,
           'xunit':'min',
           'location':0.5,
           'time_window':[['full',
                          -20,
                          +80]],
           'xlabel':'Time',
           'legend':False,
           'ylabel':'Concentration'}
    ,'use':{'title':'Glutammate release',
           'varnames':[['U_SE','k']],
           'hdf5_section':all_spines,
           'xunit':'min',
           'yunit':'\%',
           'moltiplier':1000,
           'location':0.5,
           'time_window':[['full',
                          -20,
                          +80]],
           'ylim':[98,162],
           'xlabel':'Time',
           'legend':False,
           'ylabel':'Norm. rel.',
           'numbers':True}
    ,'bdnf':{'title':'Released BDNF',
             'varnames':[['proBDNF','k'],['mBDNF','r'],['PC','b']],
             'hdf5_section':all_spines,
             'xunit':'min',
             'location':0.5,
             'time_window':[['full',
                            -20,
                            +80]],
             'xlabel':'Time',
             'legend':False,
             'thresholds':[spine_point_processes[2]['parameters']['theta_gAMPA']],
             'ylabel':'Concentration'}
    ,'fullbdnf':{'title':'Post synaptic signaling',
           'varnames':[['mBDNF','r'],['TrkB','g'],['intracell_signaling','y'],['g_factor','c']],
           # 'varnames':[['intracell_signaling','y'],['g_factor','c']],
           # 'varnames':[['g_factor','c']],
           # 'varnames':[['mBDNF','r'],['TrkB','g'],['g_factor','c']],
           'hdf5_section':all_spines,
           'xunit':'min',
           'location':0.5,
           'time_window':[['full',
                          -20,
                          +80]],
           'xlabel':'Time',
           # 'ylim':(0,40e-3),
           'legend':False,
           'ylabel':'Concentration'}
    ,'trkb':{'title':'Postsynaptic TrkB',
           'varnames':[['TrkB','g']],
           'hdf5_section':all_spines,
           'xunit':'min',
           'location':0.5,
           'time_window':[['full',
                          -20,
                          +80]],
           'xlabel':'Time',
           'legend':False,
           'ylabel':'Concentration'}
    ,'gfactor':{'title':'Effective AMPA cond.',
           'varnames':[['g_factor','c']],
           'hdf5_section':all_spines,
           'xunit':'min',
           'yunit':'\%',
           'moltiplier':100,
           'location':0.5,
           'time_window':[['full',
                          -20,
                          +80]],
           'ylim':[98,230],
           'legend':False,
           'xlabel':'Time',
           'ylabel':'Norm. cond.'}
    # ,'Peak_EPSP':{'title':'EPSP Peak',
    #        'varnames':[['v','Blues']],
    #        'hdf5_section':['pyr2005/branch_base'],
    #        'location':0.5,
    #        'time_window':['first_test_post',#'first_induction',
    #                       -60e3*10,
    #                       'end'],
    #        'xunit':'min',
    #        'xlabel':'Time',
    #        'ylabel':'%',
    #        'panel':(4,3,2)}
    ,'syn_eff':{'title':'Synaptic efficacy',
           'varnames':[['U_SE_factor','b'],['g_factor','g']],
           'hdf5_section':['Spine_0/head_0',
                           'Spine_1/head_1',
                           'Spine_2/head_2',
                           'Spine_3/head_3'],
           'location':0.5,
           'time_window':[['first_test_post',#'first_induction',
                          -20e3,
                          'end']],
           'xunit':'min',
           'xlabel':'Time',
           'legend':False,
           'ylabel':'Efficacy',
           'panel':(4,3,5)}
  ,'control':{'title':'NMDA current in control',
           'varnames':[['iampa','b'],['inmda','g']],
           'hdf5_section':['Spine_9/head_9'],
           'location':0.5,
           'time_window':[['second_test_pre',
                          -20,
                          +200]],
           'xunit':'ms',
           'legend':False,
           'xlabel':'Time',
           'ylabel':'Current'}
  ,'post_expression':{'title':'Syn current potentiated',
           'varnames':[['iampa','Blues'],['inmda','Reds']],
           'hdf5_section':['Spine_9/head_9'],
           'location':0.5,
           'time_window':[['first_test_post',
                          -20,
                          +200]],
           'xunit':'ms',
           'legend':False,
           'xlabel':'Time',
           'ylabel':'Current'}
    }

    
protocols = {
    'LTP14':{
        'BPAP_dep_stimulus_duration':2.5, # ms        
        'BPAP_hyp_stimulus_duration':0, # ms
        'n_BPAP':4,
        'activate_LTP_protocol':True,
        'nstim':25,
        'repeat_protocol':1,
        'color':'r'}
    # ,'LTP14_2':{
    #     'BPAP_dep_stimulus_duration':2.5, # ms        
    #     'BPAP_hyp_stimulus_duration':0, # ms
    #     'n_BPAP':4,
    #     'activate_LTP_protocol':True,
    #     'nstim':25,
    #     'repeat_protocol':1,
    #     'color':'r'}
    # ,'1EPSP':{
    #     'BPAP_dep_stimulus_duration':0, # ms        
    #     'BPAP_hyp_stimulus_duration':0, # ms        
    #     'activate_LTP_protocol':True,
    #     'n_BPAP':0,
    #     'nstim':70,
    #     'repeat_protocol':1,
    #     'color':'#FF4500'}
    ,'LTP11':{
        'BPAP_dep_stimulus_duration':2.5, # ms        
        'BPAP_hyp_stimulus_duration':0, # ms        
        'n_BPAP':1,
        'activate_LTP_protocol':True,
        'nstim':70,
        'repeat_protocol':1,
        'color':'b'}
     # ,'LTP14_RMBLK':{
     #     'BPAP_dep_stimulus_duration':2.5, # ms        
     #     'BPAP_hyp_stimulus_duration':0, # ms
     #     'n_BPAP':4,
     #     'activate_LTP_protocol':True,
     #     'nstim':25,
     #     'Block_RMBLK':True,
     #     'repeat_protocol':1,
     #     'color':'r'}
    # ,'LTP12':{
    #     'BPAP_dep_stimulus_duration':2.5, # ms        
    #     'BPAP_hyp_stimulus_duration':0, # ms        
    #     'n_BPAP':2,
    #     'activate_LTP_protocol':True,
    #     'nstim':50,
    #     'repeat_protocol':1,
    #     'color':'g'}
    # ,'4BPAPs':{
    #     'BPAP_dep_stimulus_duration':2.5, # ms        
    #     'BPAP_hyp_stimulus_duration':0, # ms        
    #     'n_BPAP':4,
    #     'nstim':25,
    #     'activate_LTP_protocol':False,
    #     'color':'r'}
    # ,'2BPAPs':{
    #     'BPAP_dep_stimulus_duration':3, # ms        
    #     'BPAP_hyp_stimulus_duration':0, # ms        
    #     'n_BPAP':2,
    #     'nstim':0,
    #     'activate_LTP_protocol':False,
    #     'color':'r'}
    # ,'1BPAP':{
    #     'BPAP_dep_stimulus_duration':4, # ms        
    #     'BPAP_hyp_stimulus_duration':0, # ms        
    #     'activate_LTP_protocol':False,
    #     'n_BPAP':1,
    #     'nstim':0,
    #     'repeat_protocol':1,
    #     'color':'k'}
          }

print(list(protocols.keys()))
    
# protocols = {'1EPSP':{
#     'BPAP_dep_stimulus_duration':0, # ms        
#     'BPAP_hyp_stimulus_duration':0, # ms        
#     'activate_LTP_protocol':True,
#     'color':'#FF4500'}
#     }
# deltaT = 10 ms  = post - pre 
# blue = LTP14
# verde = LTP11
# rossa = solo 4 BPAPs, calcio da VGCC
# nera = solo 1 BPAP, calcio da VGCC
# arancio = solo 1 EPSP, calcio da NMDA

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