. 2021 Jun 10;16(6):e0252291. doi: 10.1371/journal.pone.0252291

Table 8. Data for spermatozoa.

Species	Geometry			Kinematics		References
Species	B	W	Flagella	U	Notes	References
(Cricket)	110 ± 10		L = 870 ± 31.6, λ ≈ 20, h = 0.9^†.		ω = 13.3 ± 3.4^†. ^†In basic suspension medium at 18.5 ± 0.5°C and η = 1.2.	[339]
(Guinea Pig)	10.86^⋆	9.68^⋆	L = 108.55 (L_midpiece = 11.5).	9.48 ± 0.40		[100, 340]
(Rabbit)	(8.06 − 8.51)	(4.59 − 4.98)	L = (46 − 49.51) (L_midpiece = 8.81), λ = 41.6 ± 4.2^†, h = 3.3 ± 0.3^†.	(101 ± 7^‡ − 272 ± 14^♢)	ω_shallow = 17 ± 0.9, ω_deep = 18 ± 0.9. New Zealand white rabbit spermatozoa at 37°C. ^†From tracings, using the values of B, W and L. ^‡Average path velocity in shallow slide (25 μm) with ampullar fluid. ^♢Head velocity in deep slide (100 μm) with ampullar fluid. Cells diluted in BO medium were also studied.	[63, 341–343]
(Rat)	20.44	2.93	L = (171.1 − 190) (L_midpiece = 63).	(71 ± 19^† − 166 ± 32^‡)	ω = 11.27 ± 3^×. ^†Straight line velocity, measured with Computer-Assisted Sperm Analysis (CASA). ^‡Curvilinear velocity measured with CASA. Average path velocity = 93 ± 29 μm s⁻¹, lateral head displacement A_h = 9.7 ± 3.1μm. Values were also obtained with manual tracking.	[97, 100, 341]
(Stallion)	(5.33 − 6.62)	(2.79 − 3.26)	L = (40.5 − 57) (L_midpiece = (8-10.5), L_endpiece = 2.5^⋆).	86.7 ± 3.8^†	^†In still fluid. The influence of the current velocities of the fluid on the absolute speed of the spermatozoa was also measured.	[344, 345]
Acipenser baeri (siberian sturgeon)				(250 − 300)^†	ω = 60. ^†Activity lasts (2 − 3) min.	[98]
Aedes (mosquito)	(7.84^† − 8.57^‡)^⋆	(4.13^† − 5.33^‡)^⋆	L > 46^⋆†♢, h ≈ 5.		ω = (3.4^♣ − 34^♠). ^† A. triseriatus. ^‡ A. aegypti. ^♢Tail was not entirely shown in picture. ^♣Large amplitude waves. ^♠Short amplitude waves.	[102, 346]
Aleochara curtula (beetle)	15.4 ± 0.44^†		L = 84.8 ± 12.81, λ = 9.9(7 − 14.5), h = 1.3(0.8 − 2.1). Helicoidal DDW.	8.4 (3.7 − 15.2)	ω = 19.2(7.1 − 39.2). ^† L_acrosome = 4.9 ± 0.2, L_nucleus = 10.5 ± 0.4.	[347]
Asterias amurensis (starfish)				259 ± 8	Ω = 2.3 ± 0.3. The authors also estimated the torque as 600 pN nm.	[348]
Bacillus (stick insect)			Characteristic large and small waves: λ_large = (20 − 30), λ_small = (6 − 12), h_large = (9 − 15), h_small = (3 − 4).	(16 − 100)	c_large = (20 − 90), c_small = (40 − 300), ω_large = (0.9 − 2.8), ω_small = (7 − 28).	[103]
Bos (bull)	8.87^†(6.77 − 10.2)	4.74^†(4.2 − 5.4)	L = (44.2 − 63.83) (L_midpiece = (9.7 − 14.8)), n_w ≈ 1, λ = (30.5^⋆ − 40), h = 8(7.1^⋆ − 11). Cells present a 3D helical or complex (with varying amplitude) flagellar beat.	97 ± 6 (40 − 160)^‡	c = (400 − 700), Ω = 8(7.14 − 9.1), ω = 20.57 ± 3.4. ^†Average of our registered values. ^‡Cells also happen to swim in circles with velocity between 20 and 100.	[19, 41, 63, 100, 101, 247, 281, 349–353]
Bufo marinus (toad)	>7.6^†	0.69	L > 21.51^†, λ ≈ 20, h = 2.88 ± 1.13.	22.12 ± 15.9 (6.9 − 49.2)	ω = 11.74 ± 3.2(6.7 − 15.3). ^†Not entirely comprised in the figure.	[354]
Campanularia flexuosa (hydroid)	3.5^†	0.81^⋆	L = 40.	(150 − 180)	^†Head and midpiece.	[355]
Carassius auratus (goldfish)	4.2 ± 0.06^×	4.3 ± 0.06^×	L = (30.3 − 52.9)^×.	109.4 ± 9.8^×	Results obtained using automated sperm morphology analysis (ASMA) and computer assisted sperm analysis (CASA). The authors investigated the effect of mercury on the motility and morphology of the spermatozoa.	[356]
Ceratitis capitata (fly)			Characteristic large and small waves: λ_large = 30, λ_small = (5 − 8), h_large = (10 − 20), h_small = (1 − 2).	16	c_large = 120, c_small = 150, ω_large = (2 − 4), ω_small = 20.	[104, 105]
Chaetopterus (annelid)	(3.4 − 8.15^†)	(1.7 − 4.56^†)	L = 36, n_w = (1.25 − 1.4), λ = (19.3 − 25.4), h = 3.8, 2D beat.	105	c = 660, ω = 26. η = 1.4. ^†Three images are superimposed, contributing to a lack of precision in measures.	[19, 349, 353, 357, 358]
Ciona (tunicate)	(4.1 − 4.74^⋆)	(1.33^⋆ − 2.4)	L = 47.5, n_w = (1.3 − 1.57), λ = (22 − 32), h = (4.3 − 4.7), 3D and 2D beating.	165	c = (1070 − 1122.5), ω = 35. At 16°C, η = 1.4.	[19, 349, 353, 357, 358]
Colobocentrotus (sea urchin)	7.17 ± 0.13^⋆	3.1 ± 0.36^⋆	L = (35.5^⋆ − 42), n_w = (1.25 − 1.5), λ = 20.9 ± 3^⋆, h = 3.94 ± 0.95^⋆. 2D beat.	(165.6 − 193.2)	ω = 46. At 23-26°C, η = 1.8. The authors also studied the movement of ATP-reactivated sperm: U = (73.6 − 83.2), ω = 32.	[19, 106, 349]
Columba livia (pigeon)	16		L = 132 ± 11.1 (L_midpiece = 98.1 ± 11.2). More complex than a helical wave, with consistent angular velocity always CW.			[99]
Coturnix coturnix var. japonica (quail)			L = 208 (L_midpiece = 161). Irregular beat, decaying towards the end of the midpiece.	(max 50)	Ω = max4 (CW). At 20-23°C.	[99]
Culex (mosquito)	(13.7^† − 14.6^⋆)	0.48^†	L > 41.2^†‡, n_w = 3.3, λ = 15.5, h = 2.6.	6.3^⋆	^† C. pipiens quinquefasciatus. ^‡Not entirely in the picture.	[19, 346, 359]
Culicoides melleus (midge)	15.7 ± 0.4		L = 173.2 ± 1.17 (L_midpiece = 6.5 ± 0.5). Characteristic large and small waves: n_wsmall = 16.8, λ_large = 54.1 ± 1.1, λ_small = 8.7, h_large = 2.1 ± 0.9, h_small = 0.8. Planar beating.	8.3	c = 80, ω = 8.2(max20). At 25-27°C, pH10.1.	[104, 105, 360]
Cyprinus carpio (carp)				140^†	ω = 53^†. Activity lasted 200 s.	[98]
Dendraster excentricus (sand dollar)				(95.75 ± 23.8^† − 241.5 ± 46.3^‡)	^†At 7.1°C. ^‡At 24.7°C.	[361, 362]
Dicentrarchus labrax (sea bass)				120^†	ω = 70. ^†During (50 − 60) s.	[98, 363]
Didelphis (opossum)			3D beat.		At 37°C, swim in pairs.	[341, 364]
Echinus microtuberculatus (sea urchin)				120		[365]
Fugu (puffer fish)				160		[363]
Gadus morhua (cod)	(1.8 − 3.6)^†	(1.5 − 2.3)^†	L = (51.5^⋆ − 90.5), λ = 21.6^⋆‡, h = 2.25^⋆‡.	(48.3 − 201.5)^♢	ω = (52 − 55). ^†Heads can be elon-gated or round shaped. ^‡After 14s activation with sea water. ^♢At 22°C, motility lasted (7 − 800) s.	[98, 363, 366, 367]
Gallus domesticus (domestic fowl)			L = 82 (L_midpiece = 4), λ = 24.6 ± 3.6^†, h = 5.9 ± 1.5^†, dextral helix.	66.5 ± 10.1^†	c = 623.6 ± 131.6^†, Ω = 14.8 ± 2.9^†, ω = 25.4 ± 4.8^†. ^†Rapid, co-ordinated motility at 23°C in standard saline medium. Cells also swam in slow, low amplitude motility.	[99]
Hemicentrotus pulcherrimus (sea urchin)				243 ± 15	Ω = 4.8 ± 0.8. The authors also estimated the torque as 900 pN nm.	[348]
Hippoglossus hippoglossus (halibut)				(150 − 180)	ω = 55. Duration of motility: 110-120 s.	[98, 363]
Homo (human)	5.1^†(4.5 − 6.11)	3.2^†(3 − 3.45)	L = (49.7 − 56.2) (L_midpiece = (4 − 7)), λ = 32.1 ± 12.7(= c/ω). 3D beat.	30.8 ± 11.1 (7 − 50)	c = 253.8 ± 76.9(91 − 499), ω = (7.9 ± 2 − 19.1 ± 2.95). U decreases 46% in cervical mucus. Success in fecundation might be directly related to forward swimming speed (U ≥ 25 μm s⁻¹) and amplitude of lateral head displacement (A_h ≥ 7.5 μm). Authors measured, for 57 ejaculates: A_h = 5.5(2 − 10) μm. ^†Average of our registered values.	[7, 63, 96, 100, 101, 281, 341, 353, 368, 369]
Littorina sitkana (sea snail)	27^⋆†	1^⋆†	L = 25.4^⋆†(L_midpiece = 16^⋆†).	185(18^‡ − 200)	Cells swim in a spiral path doing 24 revolutions per second. ^†From illustration, obtained after superimposing two frames from a film. ^‡Backward swimming (tail first), which is more frequent in viscous fluids. U(η) is available.	[370]
Lygaeus (milkweed bug)	(4.8 − 5.24^⋆)	(0.7^⋆ − 1)	L > 29.8^†, n_w = 2.3, λ = (13 − 14.5^⋆), h = (1.3^⋆ − 2.1).		ω = (110 − 130). ^†Not entirely in the picture and estimating the end of the head and beginning of the tail.	[19, 359, 360]
Lytechinus (sea urchin)	(5.1 − 7.55^⋆)	(2.9 − 2.97^⋆)	L = 43.5, n_w = 1.45, Λ = 29.9, λ = 24.8^† (22.6 − 30), h = (4.6 − 4.7). 2D beat.	158	c = (854 − 900), ω = 30. At 16°C, η = 1.4. ^†Average of all registered values.	[19, 349, 353, 357, 358]
Megaselia scalaris (fly)	18.7 ± 0.54	0.16 ± 0.01	L = 128.7 ± 4.09. Characteristic large and small waves: n_wlarge ≈ 1.1, λ_large = (68 − 75^⋆), λ_small = 7, h_large = (9.3 − 10.27^⋆), h_small = 0.5.	117.6 ± 29.6^†	ω_large = 3.1. ^†For straight cells as they move in natural fluid (rounded and linear cells could be observed). Rounded cells moved with U = 12.7 ± 6. Cells in methyl cellulose tended to be linear and move at U = 35.5 ± 10.3.	[105, 371, 372]
Merluccius merluccius (hake)	(2.7 − 3.9)^⋆	3.1 ± 0.5	L = (30 − 50) (L_midpiece = (2.2-2.6)^⋆), n_w = (0.5 − 4), λ = (12.1 − 20.9)^†, h = (3 − 8)^‡.	(57 − 130)	[98]: ω = (56 − 57). Motility lasted for (4 − 500) s. ^†The wavelength decreases linearly with the period of swimming from λ(6 s) = 20.9 μm to λ(28.3 s) = 12.1 μm. ^‡The amplitude h remains approximately constant = 8 μm between 6 s to 17 s of activity and then decreases linearly to 3 μm at 28.3 s. [373] measured, after actvation with sea water, ω = 53, λ = 12, h = 4 and U = 82 ± 25. They also have results for 90 s and 180 s after activation and ω in function of temperature. [374] reports U = (69 − 102) and has values for the amplitude of lateral head displacement.	[98, 366, 373, 374]
Mesocricetus (hamster)	(13.8 − 15.2)	(2.51 − 3)	L = (176.5^⋆ − 250) (L_midpiece = 50.5). 3D beat.	6.75 ± 0.15	ω = 7.75 ± 1.6.	[19, 63, 100, 340, 341]
Monodelphis domestica (opossum)	17.65^⋆	8.77^⋆	L > 237.94^⋆(L_midpiece = 10.36).	(247 ± 14^† − 342 ± 34^‡)^♢	^†For single spermatozoon at 37°C in Minimum Essencial Medium (MEM). ^‡For paired spermatozoa at 37°C in MEM. ^♢Straight line velocity measured with Computer-Aided Semen Analysis (values obtained with sperm tracking are also available and values of curvilinear velocity). Lateral head displacement for paired spermatozoa A_h = 5.6 ± 2.1 μm and A_h = 11.4 ± 2.6 μm for single spermatozoa, at 37°C. Increased viscosity reduces straight line velocity for both paired and single spermatozoa, but paired spermatozoa are able to have net displacement whereas single ones moved in tight circles with poor straight line velocity.	[27]
Mus (mouse)	(7.24 − 9.44)	(3.2 − 4.48)	L = (113.4 − 134) (L_midpiece = (18.4–26.8^⋆)), n_w = 1.2, λ = (50 − 65), h = 15. 3D beat.		ω = 13.2 ± 2.5.	[19, 63, 100, 341, 353]
Myzostomus (worm)	30.8 ± 4.55		L = 52 ± 2.5. With 9+ 0 axoneme, spermatozoa can swim either with Flagellum foremost (BDW) or Head foremost (DDW) (see superscript): λ^F = 3.3 ± 2.1, λ^H = 28 ± 3, h^F = 0.9 ± 0.6, h^H = 1.7 ± 0.6. The form of the helicoidal body also changes according to the configuration of swimming: $λ_{B}^{F} = 16.3 \pm 2.6$ , $λ_{B}^{H} = 16.7 \pm 2.4$ , $h_{B}^{F} = (1 \pm 0.4 - 3.3 \pm 0.9)$ , $h_{B}^{H} = (1.4 \pm 0.5 - 3.2 \pm 0.7)$ .	(20.7 ± 9.8^† − 45.4 ± 18.3^‡)	Ω^F = 19.8 ± 5.5, ω^F = 17.5 ± 3.5, Ω^H = 20.9 ± 4.4, ω^H = 18.3 ± 2.9. M. cirriferum Leuckart observed at 15 − 21°C in seawater. ^†Flagellum foremost. ^‡Head foremost.	[375]
Oikopleura dioica (tunicate)	1	1	L = 28 (L_midpiece = 3).	75.61 ± 1.90 (max109.88 ± 1.65)^†	^†The authors examined the motile behaviour in a gradient of sperm attractant.	[376, 377]
Oncorhynchus mykiss (trout)				220^†	ω = 55. ^†Activity lasted 30 s.	[98]
Ostrea (oyster)	2.6	2.8	L = 47, λ = 25.6, h = 4.7. 2D and 3D beat.	(163.8 − 169)	ω = 43. At 23°C.	[19, 353, 354]
Ovis (ram)	10.6	6.2	L = 59, λ = 36.5, h = 7.3. 2D and 3D beat.	(132.3 − 136)	ω = 29. At 35.5°C.	[19, 353, 354]
Periplaneta americana (cockroach)	14.85^⋆†	0.95^⋆	L = 57.75^⋆, h = 6.43^⋆	(16.1 ± 1.22^‡ − 53.6 ± 3.1^♢)	^†The acrosome measured 2.08. ^‡At 15 − 16.6°C. ^♢At 37 − 39°C	[23, 346, 360]
Polyodon spathula (paddlefish)				175	ω = 50. Activity lasted 50 s.	[98]
Psammechinus (sea urchin)	1		L = (40 − 45), n_w = 1.25, λ = 24, h = 4. 2D beat.	(180 − 191.4)	c = (800 − 1000), ω = 35(30 − 40), Ω = 3.	[59, 281, 285, 353, 365, 378]
Psetta maxima (turbot)			λ = (6.7 − 10.87)^×†, h = (0.5 − 5.33)^×‡.	220	ω = 60. Motility during 200 s, with varying λ and h. ^†λ between 10.15 and 10.87 μm up to 50 s after activation and then decreases linearly to λ(142s) = 6.7 μm. ^‡The amplitude decreases almost linearly from 5.33 μm at 10.45 s to 0.5 μm at 142 s.	[98, 363]
Salmo salar (atlantic salmon)	(3.6 − 5.5)^×		L = (28.2 − 35.7)^×.	(18 − 127)	Longevity varied between 18 and 72 s. The authors observed that males with longer sperm had shorter-lived gametes.	[379]
Salmo trutta fario (trout)				(160 − 164)^†	^†At 12.5 − 16°C, 4 s after activation by fresh water. U(8 s) = (85 − 91), U(16 s) = (24 − 33) and U(26 s) = (2 − 5).	[380]
Scaphirhynchus platorynchus (shovelnose sturgeon)				200^†	ω = (48 − 50). ^†Activity lasted 48 − 50 s.	[98]
Silurus glanis (wels catfish)				130	ω = 35. Activity lasted 90 s.	[98]
Strongylocentrotus purpuratus (sea urchin)	5.6^⋆	2.5^⋆	L = (38 − 41.7^⋆), n_w = (1 − 1.5), λ = 27.7 ± 2^†, h = (4 − 4.5)^⋆.	145.3^⋆	ω = 31.1 ± 0.7^†. ^†At 18°C and η = 1.1. Values of λ and ω in function of η are available. [349] reports λ = (30 − 31.6) and ω = (25 − 31) at 16°C.	[349, 381, 382]
Sturnus vulgaris (starling)	10.3		L = 73.4.	110^†(max200)	Cells swam in three different ways: “Twist-drill” motility (TD, large majority of sperm., spin frequency and swimming velocity rose exponentially with temperature). Spin frequency = 42^† (max90) s⁻¹; “Wave” motility (Ω > 30, U > U_TD, helical flagellar wave with frequency = (3 − 10) s⁻¹; “Speedometer-cable” motility. ^†At 20°C, body temperature.	[99]
Taeniopygia guttata (zebra finch)	11.3 ± 1		L = 64.1 ± 5.7.		“Twist-drill” motility (see Sturnus vulgaris).	[99]
Tenebrio (mealworm beetle)	6.2	1.7	n_w = 4. Characteristic large and small waves: λ_large = (20 − 30), λ_small = (6 − 12), h_large = (9 − 15), h_small = (3 − 4). 2D beat.	(16 − 100)	c_large = (20 − 90), c_small = (40 − 300), ω_large = (0.9 − 2.8), ω_small = (7 − 28).	[19, 103, 104, 359]
Tripneustes (sea urchin)					ω = 60. At 25°C.	[349]
Tuhunnus thynnus (tuna)	2.3^⋆	1.13^⋆	L = 36.3^⋆, n_w ≈ 2, λ = 14.83^⋆, h = 1.67^⋆.	(215 − 340^⋆)	c = (850 − 960)^†, ω = (57 − 65)^‡. Activity lasted 140 s. ^†Apparent c = 624.3^⋆; the values were hence obtained by adding the swimming speed. ^‡Using the obtained values of c and λ (The values are in the margin of error given by one of the articles: ω = (50 − 70)).	[98, 363]