Table 2. Variables, intermediate functions, and reference parameters values.
Symbol | Description | Units | Reference value | Source |
---|---|---|---|---|
Upstream module: SAP Receptor and cGMP signalling | ||||
SAP and receptor | ||||
S | Extracellular SAP concentration | nM | Variable a | Eqs 8–11 b |
RF | Number of free SAP receptors per cell (flagellum) | cell−1 | Variable | Eqs 8–11 |
RH | Number of high activity SAP-receptor complexes per cell (flagellum) | cell−1 | Variable | Eqs 8–11 |
RL | Number of low activity SAP-receptor complexes per cell (flagellum) | cell−1 | Variable | Eqs 8–11 |
RT | Total number of SAP receptors per cell (flagellum) | cell−1 | 7.36 × 105 | F [22], S1 Fig |
θR | Receptor-bound to soluble SAP concentration conversion factor | nM cell | 6.14 × 10−5 | D (sec. 4.2.1) |
s | Sperm density in measurement chamber | cell mL−1 | 3.706 × 107 | F [22], S1 Fig |
r1 | SAP-receptor association rate constant | nM−1 s−1 | 2.70 × 10−2 | [50] |
r2 | RH to RL conversion rate constant | s−1 | 8.70 | F [22], S1 Fig |
r3 | RL inactivation rate constant | s−1 | 5.70 × 10−2 | F [22], S1 Fig |
cGMP | ||||
G | Intraflagellar cGMP concentration | nM | Variable | Eq 4 |
Gr | Resting cGMP concentration | nM | 1.25 | F [22] |
σG | Receptor-independent cGMP synthesis rate | nM s−1 | 23.65 | F [22], S1 Fig |
δG | cGMP turnover rate constant | s−1 | 18.92 | F [22], S1 Fig |
ρG | Receptor-dependent cGMP rate | cell−1 s−1 | Function | Eq 14 |
kL | cGMP synthesis rate constant per RL | s−1 | 3.25 | F [22], S1 Fig |
kH | cGMP synthesis rate constant per RH | s−1 | 40.20 | F [22], S1 Fig |
BG | Flagellum cGMP buffering power | cell−1 | 5.49 × 103 | F [10] |
θG | Conversion factor to turn cGMP molecule number into effective intraflagellar concentration | nmcell | 1.89 × 10−4 | D (sec. 4.1.2) |
KCNG channels | ||||
Fraction of open channels | Variable | Eq 15 | ||
αkn | Opening rate constant | nM−1 s−1 | 10 | F [10] |
βkn | Closing rate constant | s−1 | 257 | [48] |
gkn | Maximal conductance density | pS μm−2 | 135.30 | Ac [10] |
Ikn | Current density | fA μm−2 | Function | Eq 18 |
spHCN channels | ||||
mhc | Fraction of open gates | Variable | Eq 19 | |
Fraction of open channels | Function | |||
Steady state fraction of open channels | Function | Eq 20 | ||
τhc | Characteristic time of gating | s | Function | Eq 21 |
h1 | Half-activation voltage | mV | -50.80 | [62, 76] |
h2 | Voltage sensitivity of activation | mV | 6.60 | [62, 76] |
h3 | τhc basal value | s | 9.18 × 10−2 | F [62], S2 Fig |
h4 | τhc amplitude | s | 1.10 | F [62], S2 Fig |
h5 | Voltage of maximum τhc | mV | -42 | F [62], S2 Fig |
h6 | Characteristic width of τhc | mV | 18.50 | F [62], S2 Fig |
ghc | Maximal conductance density | pS μm−2 | 193.50 | Ad [10] |
Ihc | Current density | fA μm−2 | Function | Eq 22 |
CaV + BK module | ||||
Equilibrium parameters for V and C | ||||
gL | Leakage conductance density | pS μm−2 | 1.94 | F [10]e |
EL | Reversal potential of leakage current | mV | 80.05 | D (Eq 3) |
σC | CaV-independent Ca2+ source rate constant | nm s−1 | 1.14 × 108 | D (sec. 4.6) |
δC | Ca2+ extrusion rate constant | s−1 | 1.16 × 106 | MF |
CaV channels | ||||
Fraction of open channels | Variable | Eq 30 | ||
Fraction of closed channels | Variable | Eq 31 | ||
αcv | Inactivation recovery rate | s−1 | Function | Eq 27 |
βcv | Opening rate | s−1 | Function | Eq 28 |
γcv | Inactivation rate | s−1 | Function | Eq 29 |
v1 | Inactivation recovery rate constant | mV−1 s−1 | 10.10 | MF |
v2 | inactivation recovery rate voltage inflexion point | mV | -55 | MF |
v3 | Inactivation recovery rate voltage sensitivity | mV | 1.42 | MF |
v4 | Opening rate constant | mV−1 s−1 | 33.80 | MF |
v5 | Opening rate voltage inflexion point | mV | -39 | MF |
v6 | Opening rate voltage sensitivity | mV | 2.10 | MF |
v7 | Inactivation rate constant | mV−1 s−1 | 8.10 | MF |
v8 | Inactivation rate voltage inflexion point | mV | -18 | MF |
v9 | Inactivation rate voltage sensitivity | mV | 7.50 | MF |
gcv | Maximal conductance density | pS μm−2 | 185.16 | MFf |
Icv | Current density | fA μm−2 | Function | Eq 33 |
BK channels | ||||
Fraction of open channels | Variable | Eq 45 | ||
αbk | Opening rate | s−1 | Function | Eq 46 |
βbk | Closing rate constant | s−1 | 97 | MF |
b1 | Opening rate constant | nM−1 s−1 | 0.97 | MF |
b2 | Opening rate [Ca2+]i inflexion point αbk | nM | 316 | MF |
b3 | Opening rate [Ca2+]i sensitivity | nM | 30 | MF |
gbk | Maximal conductance density | pS μm−2 | 214.50 | MFg |
Ibk | Current density | fA μm−2 | Function | Eq 47 |
CatSper + NHE module | ||||
Equilibrium parameters for V and C | ||||
gL | Leakage conductance density | pS μm−2 | 9 | MFh |
EL | Reversal potential of leakage current | mV | -20.4 | D (Eq 3) |
σC | CatSper-independent Ca2+ source rate constant | nM s−1 | 3.72 × 107 | D (sec. 4.6) |
δC | Ca2+ extrusion rate constant | s−1 | 4.27 × 105 | MF |
NHE exchangers | ||||
Fraction of active exchangers | Variable | Eq 23 | ||
αnh | Activation rate | s−1 | Function | Eq 24 |
βnh | Inactivation rate | s−1 | Function | Eq 25 |
n1 | Activation rate constant | mV−1 s−1 | 1.51 | MF |
n2 | activation rate voltage inflexion point | mV | -65 | MF |
n3 | activation rate voltage sensitivity | mV | 7 | MF |
n4 | inactivation rate constant | mV−1 s−1 | 0.90 | MF |
n5 | inactivation rate voltage inflexion point | mV | -30 | MF |
n6 | inactivation rate voltage sensitivity | mV | 7 | MF |
Jmax | Maximal H+ flux by total active exchangers | fmol s−1 | 3 × 10−7 | MF |
KNa | Na+ dissociation constant | mM | 25 | MF [69]i |
CatSper channels | ||||
Fraction of open channels | Function | Eq 34 | ||
mcs | Fraction of channels with voltage- and pH-dependent gate open | Variable | Eq 35 | |
hcs | Fraction of channels with Ca2+-dependent gate not inactivated | Variable | Eq 36 | |
vcs | pH-dependent voltage leading to half-maximal gate opening at equilibrium | mV | Function | Eq 40 |
s1 | Voltage-dependent gate opening rate constant | s−1 | 270 | MF |
s2 | mcs voltage sensitivity | mV | 10 | MF [12, 83], S3 Fig |
s3 | Amplitude of vcs response to pHi | mV | 79.70 | F [72], S3 Fig |
s4 | pHi leading to half-maximal vcs value | 7.20 | MF [72], S3 Fig | |
s5 | Hill-coefficient of vcs | 35.52 | F [72] | |
s6 | pHi-independent, minimal vcs value | mV | -41 | MF, S3 Fig |
s7 | Scaling factor | nM | 1 | MF |
s8 | [Ca2+]i leading to half-maximal hcs inactivation (IC50) | nM | 700 | MF |
s9 | Hill coefficient of Ca2+-inactivation curve | 2.50 | MF | |
s10 | Ca2+-inactivation rate constant | nM−1 s−1 | 1.40 × 10−4 | MF |
gcs | Maximal conductance density | pS μm−2 | 39.90 | MF j [72] |
Ics | Current density | fA μm−2 | Function | Eq 44 |
Common variables and parameters | ||||
V | Membrane potential | mV | Variable | Eq 1 |
Em | Resting membrane potential | mV | -40 | [80, 84, 85, 10] |
Cm | Specific capacitance of flagellum membrane | pF μm−2 | 1 × 10−2 | [86] k |
vf | Intraflagellar volume | fL | 1.60 | [48] |
sf | Surface area of flagellar membrane | μm2 | 30 | D l |
[K+]o | Extracellular potassium concentration | mM | 9 | [87] m |
[K+]i | Resting intraflagellar potassium concentration | mM | 219 | [66] |
[Na+]o | Extracellular sodium concentration | mM | 423 | [87] m |
[Na+]i | Resting intraflagellar sodium concentration | mM | 20 | [81] |
pHo | Extracellular pH | 8 | [87] m | |
pHr | Resting intraflagellar pH | 7.10 | [81] | |
[Ca2+]o | Extracellular calcium concentration | mM | 10 | [87] m |
Cr | Resting intraflagellar calcium concentration | nM | 100 | [81, 14] |
C | Intraflagellar calcium concentration | nM | Variable | Eq 6 |
H | Intraflagellar proton concentration | M | Variable | Eq 5 |
σH | NHE-independent proton source rate constant | M s−1 | 1.50 × 10−7 | D (sec. 4.6) |
δH | Proton extrusion/consumption rate constant | s−1 | 1.28 | D (Eq 5) |
EK | K+ equilibrium potential | mV | -80 | Dn |
Ehc | Reversal potential of spHCN current | mV | -18 | [61, 62, 63] |
ECa | Ca2+ equilibrium potential | mV | 144 | Do |
a The column entries are a numeric value if the quantity is a parameter or a label whether it is a variable or a function
b The column entries: are a bibliographic reference unlabelled if the values were used as reported in the original publication or with labels F and MF if further processed; a derivation labelled D followed by a reference to the section where its derivation is explained; or a reference to the defining equation(s) defining a variable or function.
c Taking into account the membrane surface of the flagellum sf, and KCNG unitary conductance (110 pS [59]), this implies ∼37 channels per flagellum
d Taking into account the membrane surface of the flagellum sf, and spHCN unitary conductance (43 pS [63]), this would imply ∼135 channels per flagellum
e Fitted in the Upstream module
f Taking into account the membrane surface of the flagellum sf, and CaV unitary conductance (4.70 pS [82]), this implies ∼1182 channels per flagellum
g Taking into account the membrane surface of the flagellum sf, and BK unitary conductance (70 pS [42]), this implies ∼92 channels per flagellum
h Fitted in the complete CatSper + NHE module to conserve the initial drop of membrane potential, whereas allowing calcium signal to return to levels close to resting value after spike-train finishes
i Based on figure 1 of [69]
j Taking into account the membrane surface of the flagellum sf, and assuming that the Ca2+ unitary conductance of CatSper might be below 1 pS, this implies ∼1200 channels per flagellum
k Conventional value for eukaryotic cytoplasmic membranes
l Calculated as the lateral surface of a cylinder with length 40 μm and diameter 0.240 μm
m Typical concentration in artificial seawater used for embryology experiments on marine organisms
n Calculated with the Nernst equation, , having temperature T = 17°C
o Calculated with the Nernst equation, setting temperature T = 17°C, and using the Cr parameter as internal concentration value