Milk polar lipids composition and functionality: a systematic review

Figures & data

Figure 1. The structure of the milk fat globule.

Table 1. Search terms.

Matrix	Component	Exclude
Milk	Phospholipid	Peptide
Bovine	Phosphatid*	Plant
Cow	Sphingo*
Human	Glycolipid*
Breast	“polar lipid”
Mother	Cerebro*
Lact*	Ganglio*

Figure 2. Database search results according the Preferred Reporting Items for Systematic Reviews (PRISMA) statement (Moher et al. 2009).

Table 2. Mean polar lipid content (mg/100 mL milk) and proportion of individual polar lipids (mg/100 mL milk and % of total polar lipid) in human milk.

Author	Region	Lactation stage	Total polar lipid mg (or % of total lipid)^a	PC	PE	PI	PS	SM	Other	Analysis	Outcome trend
				mg [or % of total polar lipids]
Preterm
J. Bitman et al. 1983	USA n = 28	Day 3 Day 7 Day 21 Day 42 Day 84	nm	–	–	–	–	–	–	TLC	Preterm milk is richer in polar lipid. Polar lipid ↓ over lactation.
Holmes-McNary et al. 1996	USA n = 17	MM: Day 27–32	–	90 μmol/L milk	–	–	–	104 μmol/L milk		TLC
Yassar Y. Al-Tamer and Mahmood 2004	Iraq n = 29	Colostrum: Day 3–5	24.48	–	–	–	–	–	–	TLC
Shoji et al. 2006	Japan n = 14 n = 18 n = 14	Colostrum: Day 1–3 TM: Day 4–7 MM: Day 30–64	20–25^b (0.9) 28^b 25.5^b (0.6)	(28.6) - (22.5)	(14.8)^b (14)^b (13.6)^b	(7)^b (6.7)^b (6.2)^b	(6)^b (5.7)^b (6.5)^b	(36) - (42.3)	LPC (9.8)^b(9.6)^b (10)^b	TLC	SM↑, PC↓ and total polar lipid↓ from colostrum to MM.
Maas et al. 2017	Germany n = 34	Day 6–85	–	nm	–	–	–	–	–	ESI-MS	PC↓ in term milk.
Lindahl et al. 2019	Ireland n = 57	Colostrum: Day <5 TM: Day 5–2 weeks MM: >2–<15 weeks	117.3 95.49 57.49	19.31 (16.5)^b 13.09 (13.7)^b 6.92 (12.0)^b	87.85 (74.9)^b 75.25 (78.8)^b 46.64 (81.1)^b	–	–	10.13 (8.6)^b 7.21 (7.6)^b 3.93 (6.8)^b	–	LC-MS	Polar lipid↓ over lactation. Total polar lipid, PC & SM ↑ in preterm milk.
Wei, Jin, et al. 2019	China n = 10	Colostrum: Day 1–7 TM: Day 8–14 MM: Day 27–33 Day 57–63 Day 87–93	27.47 25.53 24.31 21.87 20.18	8.42 7.48 6.64 6.06 5.54	7.72 7.44 7.35 5.83 5.32	0.92 0.85 0.81 0.66 0.51	1.25 1.14 1.09 0.82 0.91	9.15 8.63 8.42 8.50 7.89	–	P-NMR	SM most abundant (27.44%–43.40% of total polar lipid), SM relatively constant in both term and preterm milk.
Dei Cas et al. 2020	Italy n = 5 (Mediterranean, carnivore or vegetarian diet)	MM: Day 10	–	–	–	–	–	nm	–	LC-MS	No difference in SM between preterm and term milk. SM not affected by diet.
W. Xu, Chen, et al. 2020	China n = 6	Colostrum: Day 2–6	nm	nm	nm	–	–	–	–	LC-MS	↑PE and PC in colostrum compared to term milk.
Shunova et al. 2020	Germany n = 21	MM Day 8–28	–	21.5 ± 4.5 mg choline/150 mL breast milk, out of which PC was 7.4 ± 2.1%	–	–	–	–	–	Tandem MS
Full-term
J. Bitman et al. 1983	USA n = 6	Day 3 Day 7 Day 21 Day 42 Day 84	nm	–	–	–	–	–	–	TLC	Preterm milk richer in PL. Polar lipids↓ over lactation.
Zeisel, Char, and Sheard 1986	nm	MM: Day 31–84		146–220 nmol/mL				107–206 nmol/mL		TLC	PC and SM ↑ in hind milk (only hind milk data presented). No diurnal variation or lactation stage effect on SM and PC concentration.
Glew et al. 1995	Nigeria n = 13 n = 3	MM: 1–4 months Moderately malnourished Severely malnourished	0.338 μmol/mL 0.372 μmol/mL	(27.7) (22.1) Range: (20–30)	(15.2) (12.8) Range: (12–15)	PI + PS: (5.9) (5.6) Range: (5)	PI + PS: (5.9) (5.6) Range: (5)	(51.3) (59.5) Range: (50–60)	–	TLC	No differences in the major polar lipid classes between the groups with moderately and severely malnourished.
Holmes-McNary et al. 1996	USA n = 16	MM: Day 27–32	–	82 μmol/L	–	–	–	124 μmol/L		TLC	Inconclusive effect of dietary choline intake on PC and SM.
Barbas and Herrera 1998	Spain n = 8	Colostrum: Day 3–5 MM: Day 21	9.69 mg/dL 8.91 mg/dL	–	–	–	–	–	–	TLC
Holmes, Snodgrass, and Iles 2000	London n = 8	Colostrum: Day 2–6 MM: Day 7–22	–	0.23 mmol/L 0.48	–	–	–	0.09 mmol/L 0.10	–	NMR	7 days considered as mature milk in this study. Data represents both preterm and term milk. ↑ PC and SM over lactation.
L. Wang et al. 2000	Japan n = 20	Week 1	–	(23.1)	(36.1)	(3.5)	(6.7)	(30.6)	–	TLC
Yassar Y. Al-Tamer and Mahmood 2004	Iraq n = 35	Colostrum: Day 3–5	29.78							TLC
Ilcol et al. 2005	Turkey n = 21 n = 95 n = 14 n = 12 n = 11	Colostrum: Day 0–2 MM: Day 12–180 Day 12–28 Day 75–90 Day 165–180	nm	146 μmol/L 104 103 155 97	–	–	–	129 μmol/L 94 91 97 84	–	TLC	PC and SM constant over lactation.
Sala-Vila et al. 2005	Spain n = 30 n = 17 n = 19	Colostrum: Day 1–5 TM: Day 6–15 MM: Day 15–30	202 nmol/mL 209 147	38.40 37.69 31.26	5.86 8.55 12.76	6.03 5.21 5.89	7.91 8.17 10.35	40.49 39.20 41.03	–	HPLC-ELSD	PS and PE ↑ over lactation; SM was constant. PC↓ in MM only.
Y. Y. Al-Tamer and Mahmood 2006	Iraq n = 25 n = 25 n = 15	MM: 4–6 months	High urban: 33.59 Suburban: 31.15 Low urban: 15.94	–	–	–	–	–	–	Colorimetric	↓ PL at 5–6 than 4–5 months. ↓ PL for women under nutritional stress.
Shoji et al. 2006	Japan n = 14 n = 15 n = 15	Colostrum: Day 1–3 TM: Day 4–7 MM: Day 30–64	(1.1) (0.5)	nm (27.2)	(14)^b (16)^b (13.8)^b	(6.1)^b (6.7)^b (8)^b	(4.3)^b (4.5)^b (6.3)^b	nm (36.0)	(10.9) ^b (9.8)^b (9.6)^b	TLC	Polar lipid ↓ over lactation. PL content (mg/dL) in TM and MM differed in term and preterm milk In both term and preterm groups, PL content (% of total lipid) of MM was lower than in colostrum.
Benoit et al. 2010	France n = 4	MM: 3–6 months	nm	(19)	(21.3)	PI + PS: (16.4)	PI + PS: (16.4)	(43.3)	–	TLC	SM ↑ in breastmilk than bovine milk.
Fischer et al. 2010	USA n = 48 n = 51	MM: Day 45 Placebo Supplement group	–	107 nmol/mL 113	–	–	–	67 nmol/mL 67		LC-MS	Dietary choline intake correlated with PC concentration. Genotype can influence the concentrations of choline and its metabolites in breast milk.
Garcia et al. 2011	France n = 5 n = 7 n = 16	Colostrum: Day 3–4 TM: Day 6–10 MM: Day 30	72 55 45	nm	nm	nm	nm	nm	–	TLC	Sample size includes both preterm (n = 14) and term milk (n = 8).
Lopez and Ménard 2011	France n = 3	MM: 2–7.5 months	1. 4.08 ± 0.21 mg/g of fat 2. 3.05 ± 0.20 3. 3.52 ± 0.63	1. (23.5 ± 0.5) 2. (20.4 ± 2.1) 3. (19.3 ± 2.4)	1. (15.1 ± 1.0) 2. (10.1 ± 1.6) 3. (14.3 ± 2.2)	1. (9.5 ± 1.1) 2. (11.7 ± 0.5) 3. (9.2 ± 0.6)	1. (15.6 ± 0.1) 2. (12.3 ± 0.8) 3. (18.4 ± 0.7)	1. (36.4 ± 1.9) 2. (45.4 ± 2.6) 3. (38.8 ± 4.5)	–	HPLC-ELSD	MM contained ∼3–4 mg polar lipid/g fat (0.3–0.4%, w/w).
Garcia et al. 2012	France n = 22	nm	213.3 μg/mL	60.3 μg/mL	41.5 μg/mL	11.1 μg/mL	22.1 μg/mL	78.3 μg/mL	LPC = 0	P-NMR	–
Giuffrida et al. 2013	Singapore n = 50	MM: 4 weeks	23.8^c mg/100 g	6.0^c (25.2)	6.8^c (28.6)	1.1^c (4.6)	1.4^c (5.9)	8.5^c (35.7)	–	HPLC-ELSD	Average 23.8 mg polar lipid/100 g in milk corresponded to an estimated mean intake of 140 mg polar lipid/day in a 4-week old exclusively breast-fed infant.
Thakkar et al. 2013	Singapore Female n = 25 Male n = 25	MM: Day 30 Female Male Combined Day 60 Female Male Combined Day 120 Female Male Combined Average	22.01 24.03 23.02 18.73 22.83 20.78 22.02 26.29 24.24 22.7^b	5.81 6.12 5.97 4.43 5.25 4.84 4.52 5.32 4.94 5.3 (23)^b	6.47 7.05 6.76 5.70 7.02 6.36 7.24 8.85 8.08 7.1 (31)^b	0.99 1.15 1.07 1.00 1.27 1.13 1.51 1.81 1.67 1.3 (6)^b	0.66 0.85 0.75 0.70 0.80 0.75 0.86 0.94 0.91 0.8 (4)^b	8.07 8.86 8.47 6.93 8.49 7.71 7.89 9.37 8.66 8.3 (37)^b	–	HPLC-ELSD	↓ PC over lactation, unlike other polar lipid classes.
X. Zou et al. 2013	Denmark n = 45	Colostrum: Day 1–5 TM: Day 6–15 MM: Day >16	4.42 5.86 5.06	1.26 (28.46) 1.50 (25.73) 1.07 (21.33)	0.41 (9.33) 0.77 (13.16) 0.76 (15.04)	0.36 (8.31) 0.40 (6.81) 0.41 (8.17)	0.56 (12.67) 0.81 (13.88) 0.84 (16.71)	1.82 (41.23) 2.37 (40.42) 1.97 (38.75)	–	HPLC-ELSD	Diameter of MFG is negatively correlated with polar lipid concentration.
Davenport et al. 2015	USA n = 15 n = 13 n = 28	5 weeks Choline intake: 480 mg/day 930 mg/day All	–	64 μmol/L 61 63	–	–	–	172 μmol/L 177 175	–	LC-MS	Choline intake had no notable effect on PC and SM concentration.
Roy et al. 2015	India n = 11 n = 8 n = 9 n = 5 n = 7 n = 9	Colostrum: Day 0–3 High income Medium income TM: Day 4–7 High income Medium income MM: >7 days High income Medium income	110.27 53.23 78.03 44.15 33.31 25.95	–	–	–	–	–	–	Spectrophotometry	Total polar lipid content ↓ over lactation and across income groups.
Claumarchirant et al. 2016	Spain (n = 65) Madrid Valencia/Murcia (VM)	Colostrum: Day 0–5 Madrid VM TM: Day 6–15 Madrid VM MM: 1 month Madrid Valencia Murcia 3 months Madrid Valencia Murcia 6 months Madrid Valencia Murcia 9 months VM 12 months VM	37.21 31.49 43.70 53.53 39.18 42.18 39.19 35.90 32.15 34.87 26.56 33.41 28.12 26.00 29.00	4.87 4.51 5.97 8.09 5.42 6.55 5.67 5.06 4.92 4.86 3.79 5.02 4.13 3.63 4.21	12.36 10.46 13.90 15.90 11.98 12.68 11.58 10.71 9.51 11.36 8.29 10.24 8.37 7.89 6.49	3.13 3.12 4.30 6.32 4.15 5.20 4.60 4.23 4.13 4.06 3.12 4.23 3.50 3.07 3.83	3.49 3.43 4.66 6.74 4.45 5.55 4.90 4.56 4.39 4.30 3.39 4.49 3.73 3.25 4.02	13.36 9.95 14.86 16.49 13.20 12.19 12.44 11.34 9.20 10.29 7.96 9.44 8.39 8.15 10.44	–	HPLC-ELSD	PI, PS and PC was ↑ in the coastal area than in the central area.
Giuffrida et al. 2016	China n = 539 (total) n = 38 n = 45 n = 30 n = 113 n = 22 n = 21 n = 38 n = 81 n = 120 n = 113 n = 112 n = 345	Colostrum: Day 0–5 Guangzhou Beijing Suzhou Average TM: Day 6–15 Guangzhou Beijing Suzhou Average MM: 0.5–8 months Guangzhou Beijing Suzhou Average	33.2 33.0 38.9 33.0 25.6 24.4 34.9 28.5 25.3 22.3 26.0 24.2	12.5 (37.7) 10.9 (33.0) 12.6 (32.4) 12.0 (36.4) 11.3 (44.1) 8.3 (34.0) 11.9 (34.1) 10.1 (35.4) 8.6 (34.0) 7.6 (34.1) 8.5 (32.7) 8.2 (33.9)	9.9 (29.8) 7.6 (23.0) 12.6 (32.4) 8.5 (25.8) 5.6 (21.9) 7.3 (29.9) 10.8 (30.9) 8.2 (28.8) 7.1 (28.1) 5.3 (23.8) 7.3 (28.1) 6.4 (26.4)	1.8 (5.4) 1.6 (4.8) 2.3 (5.9) 1.8 (5.5) 1.2 (4.7) 1.5 (6.1) 2.4 (6.9) 1.8 (6.3) 1.5 (5.9) 1.2 (5.4) 1.7 (6.5) 1.5 (6.2)	1.3 (3.9) 1.8 (5.5) 1.7 (4.4) 1.5 (4.5) 0.8 (3.1) 1.0 (4.1) 1.3 (3.7) 1.1 (3.9) 1.0 (4.0) 0.9 (4.0) 1.2 (4.6) 1.0 (4.1)	7.7 (23.1) 10.9 (33.0) 9.7 (24.9) 9.1 (27.6) 6.8 (26.6) 6.2 (25.4) 8.5 (24.4) 7.3 (25.6) 7.1 (28.1) 7.3 (32.7) 7.4 (28.5) 7.2 (29.8)	–	HPLC-ELSD	Average PL of the 3 cities and total PL ↓ over lactation PL distribution (%) was similar across lactation stages. Highest total polar lipid content was observed in human milk from Suzhou.
Yao et al. 2016	China n = 10	MM: 16–30 days	nm	(24.4)	(25.3)	(7.9)	(13.1)	(29.3)	–	HPLC-ELSD
Ma et al. 2017	Malaysia n = 48	Colostrum TM: MM: 2 months 6 months 12 months	352.4 mg/L 273.0 mg/L 147.1 mg/L 187.5 mg/L 204.7 mg/L	76.7 48.6 21.0 23.7 27.6	89.9 100.0 39.3 66.1 78.4	16.6 9.6 6.3 5.9 6.5	125.8 90.9 14.8 15.5 16.2	39.7 20.9 57.4 70.4 71.5	Lyso-PC 2.5; 2.1; 5.5; 1.3; 1.0 Lyso-PE 1.2; 0.9; 2.8; 4.5; 3.5	HPLC-MS	Strong correlation between total polar lipid concentration and fat concentration.
Maas et al. 2017	Germany n = 9	Day 6–118	nm	nm	–	–	–	–	–	ESI-MS	PC ↑ in preterm milk versus term milk.
K. Li et al. 2018	China n = 30	Colostrum: Day 1 TM: Day 14 MM: Day 42	nm	nm (↑ over lactation)	nm	nm	nm (↑ over lactation)	nm (↑ over lactation)	–	UPLC-QTOFMS	Variations in PL and SL over lactation were independent of dietary intake.
Tavazzi et al. 2018	Singapore n = 50	MM: Day 30–120	nm	5.39	2.85	1.82	nm	9.28	–	HPLC-MS
Lindahl et al. 2019	Ireland n = 22	Colostrum: Day <5 TM: Day 5-<2 weeks MM: 2-<15 weeks	67.74 48.65 36.94	11.44 (16.9)^b 6.56 (13.5)^b 4.50 (12.2)^b	49.40 (72.9)^b 37.86 (77.8)^b 29.15 (78.9)^b	–	–	6.90 (10.2)^b 4.23 (8.7)^b 3.29 (8.9)^b	–	LC-MS	Total polar lipid concentration ↓ over lactation.
McJarrow et al. 2019	United Arab Emirates TM n = 41 MM n = 40	TM: Day 5–15 MM: 6 months	26.9 22.0	6.6 (25) 3.0 (14)	6.6 (25) 8.0 (36)	1.1 (4) 0.7 (3)	2.9 (7) 1.6 (7)	9.1 (34) 8.3 (38)	5.4 (2) 4.0 (1.8)	HPLC-MS
Wei, Jin, et al. 2019	China n = 10	Colostrum: Day 1–7 TM: Day 8–14 MM: Day 27–33 Day 57–63 Day 87–93	25.83 24.78 23.67 23.63 22.57	7.55 7.21 6.74 6.06 5.69	7.39 7.07 6.92 6.89 7.13	1.05 0.93 0.64 1.00 0.75	1.29 1.21 1.03 1.15 0.85	8.54 8.37 8.34 8.54 8.15	–	P-NMR	Concentration of SM remained constant in term and preterm milk.
Dei Cas et al. 2020	Italy n = 18	MM: Day 10	–	–	–	–	-	nm	–	LC-MS	No difference in SM content between preterm and term milk. SM showed very low variability (not influenced) between the 3 diet types. SM in diet was not quantified but differed based on food sources.
B. Y. B. Y. Fong, Ma, and MacGibbon 2020	China n = 20	MM: 0.5 month 1 month 2 months 3 months 4 months 6 months 8 months Average	21.3 19.3 19.5 23.6 20.7 24.2 24.6 21.9	5.25 (25) 4.67 (24) 4.29 (22) 5.11 (22) 4.06 (20) 4.74 (20) 5.17 (21) 4.75 (22)	6.68 (31) 6.51 (34) 7.62 (39) 8.63 (37) 8.88 (43) 9.07 (37) 9.50 (39) 8.13 (37)	0.70 (3) 0.56 (3) 0.58 (3) 0.69 (3) 0.86 (4) 0.94 (4) 0.92 (4) 0.75 (3.4)	0.93 (4) 0.67 (3) 0.71 (4) 0.86 (4) 0.79 (4) 1.50 (6) 1.08 (4) 0.94 (4)	7.35 (35) 6.54 (34) 5.97 (31) 7.81 (33) 5.79 (28) 7.52 (31) 7.57 (31) 6.94 (32)	LPE + LPC 3.5 (2) 3.4 (2) 3.0 (1) 4.9 (1) 3.4 (1) 4.0 (2) 3.4 (1) 3.7 (1.6)	HPLC-MS	SM, PC & PS concentration remained relatively constant throughout the 7 timepoints but total PL concentration ↑.
Selvalatchmanan et al. 2021	Singapore n = 20	MM: Day 62–101	nm	nm	nm	nm	nm	nm	nm	LC-MS	PE concentration showed the strongest morning-evening variation. PI, PS and PC ↑ in evening milk. SM did not show diurnal variation.
Reported range^d		Colostrum MM	25.8–38.9 14.7–26.0	7.6–12.6 2.1–8.6	7.4–12.6 2.9–9.5	1.1–2.3 0.6–1.8	1.3–12.6 0.7–1.6	4.0–10.9 5.7–9.3

HPLC: high-performance liquid chromatography; ELSD: evaporative light scattering detection; P-NMR: phospho-nuclear magnetic resonance spectroscopy;               TLC: thin layer chromatography; nm: not mentioned; TM: transitional milk; MM: mature milk

^aTotal polar lipid content refers to the cumulation of PC, PE, PI, PS and SM only (5 major polar lipid classes)

^bValue calculated/interpreted based on data provided by author

^cValue represents an average of the dataset

^dConsidering full-breast studies only described/calculated in mg/100 ml (or mg/100g), including different regions and not differentiation for detection methods.

Table 3. Mean polar lipid content (mg/g of fat) and proportion of individual polar lipids (mg and % of total polar lipid) in bovine milk.

Author	Species/region	Lactation stage	Total polar lipid mg (or % of total lipid)^a	PC	PE	PI	PS	SM	Analysis	Outcome trend
				mg (or % of total polar lipid)
Christie, Noble, and Davies 1987			(22.78 mg/100 g milk)	(27.56)	(37.06)	(6.76)	(3.05)	(25.57)	HPLC
Joel Bitman and Wood 1990	Holstein n = 12	Colostrum: Day 3 TM: Day 7 MM: Day 42 Day 180	(20.4 mg/100 ml milk) (35.4) (25.1) (12.8)	5.8 (28.0) 8.9 (25.1) 6.7 (26.4) 4.5 (35.1)	6.4 (31.0) 10.0 (27.8) 7.9 (31.1) 2.6 (19.8)	0.8 (4.1) 1.6 (4.6) 1.3 (5.2) 1.5 (11.8)	1.6 (8.1) 3.0 (8.4) 2.1 (8.5) 0.3 (1.9)	5.8 (28.7) 11.9 (34.1) 7.1 (28.7) 3.9 (31.4)	TLC	Total polar lipid content ↓ as lactation progressed. % PC ↑, PE and PS ↓ at 180 days, SM remained constant and PI ↑ at 180 days.
Sharma, Sachdeva, and Singh 2000	Holstein Fresiane x Red Dane x Sahiwal (cross-bred) India n = 9 (3 of each lactation stage)	Early: Day 45) Mid: Day 125 Late: Day 280	(0.92) (1.02) (1.2)	0.21% of lipids 0.31% 0.23%	0.28% 0.23% 0.38%	0.01% 0.10% 0.10%	0.17% 0.14% 0.12%	0.25% 0.27% 0.36%	TLC	Total polar lipid, PE and SM concentration ↑ in late lactation.
Avalli and Contarini 2005	Bulk milk tank Italy		8.87 (0.031)	2.98 (33.5)	2.92 (32.9)	0.55 (6.2)	0.26 (2.9)	2.17 (24.5)	HPLC-ESLD
Rombaut, Van Camp, and Dewettinck 2006	De-identified sample Belgium		9.42 (0.038)	10.4 (27.3)	12.6 (33.1)	1.96 (5.2)	3.52 (9.3)	9.52 (25)	HPLC-ESLD
Rodríguez-Alcalá and Fontecha 2010	Breed unknown Spain	nm	(0.36)	(25.92)	(38.55)	(6.5)	(7.67)	(21.34)	HPLC-ELSD
Garcia et al. 2012	Breed unknown France n = 15	nm	197.1	60.3 (30.6)	61.8 µg/mL (31.4)	7.6 (3.9)	21.3 (10.8)	46.1 (23.4)	P-NMR
Argov-Argaman et al. 2013	Holstein-Friesian Netherlands n = 204	First lactation: Day 67–263	AA: (1.22) KK: (0.94)	AA: (29.99) KK: (30.06)	AA: (9.10) KK: (8.69)	AA: (17.64) KK: (17.11)	AA: (19.73) KK: (19.35)	AA: (23.54) KK: (24.80)	HPLC-ELSD
Kiełbowicz, Micek, and Wawrzeńczyk 2013	Unidentified Poland n = 4		7.39^b^c (0.026)	11.68^b^c (45.3)	8.47^b^c (32.6)	2.42^b^c (9.3)	2.15^b^c (8.3)	1.15^b^c (4.4)	LC-CAD
Walker et al. 2013	Australia Cows calved in Autumn n = 16 Spring n = 16	MM	6.39 6.44	2.03 (31.8) 2.05 (31.8)	2.42 (37.9) 2.47 (38.4)	–	–	1.94 (3.0) 1.92 (3.0)	HPLC	PE and PC were ↑ in late lactation in both autumn and spring groups. Polar lipid concentration ↑ over lactation while MFG size ↓ which can confound the effect of season on polar lipid concentration.
Artegoitia et al. 2014	Holstein USA n = 12 n = 12 n = 10	Early: Week 1 Week 2 Week 3 Mid: Week 7 Week 10 Week 13 Late: Week 31 Week 34 Week 37	–	μmol/L 170 149 167 202 178 183 582 701 694	–	–	–	μmol/L 52.8 44.9 45.9 33.8 30.7 27.8 35.8 36.7 36.7	HILIC LC-MS	PC in late lactation ↑ than PC reported in human milk literature over lactation. SM in late lactation ↓ than found in human milk.
Contarini et al. 2014	Holstein Italy n = 30 n = 30 n = 10 n = 10 n = 10 n = 10 n = 10 n = 10	Multiparous Primiparous 24 hours 48 hours 72 hours 96 hours 120 hours 5 months	g/100 g milk 0.02 0.02 0.04 0.02 0.02 0.02 0.02 0.02	(26.3) (26.5) (27.1) (26.0) (25.4) (25.8) (25.8) (28.4)	(28.9) (27.3) (23.4) (24.7) (26.8) (27.1) (27.8) (38.7)	(7.6) (7.9) (6.0) (8.3) (7.1) (7.6) (7.9) (9.6)	(5.9) (9.8) (7.5) (9.3) (8.9) (8.0) (8.8) (4.6)	(31.2) (28.4) (36.0) (31.6) (31.7) (31.4) (29.6) (18.7)	HPLC-ESLD	Parity did not affect total polar lipid concentration.
Zheng, Jiménez-Flores, and Everett 2014	Jersey New Zealand	Mid n = 9 n = 18 n = 9	M1: 4.69 M2: 4.78 M3: 5.00	(26.94) (27.35) (27.40)	(27.15) (27.30) (27.35)	(5.84) (5.77) (5.73)	(14.12) (14.55) (15.02)	(25.62) (25.59) (25.45)	HPLC
Cengiz et al. 2016	Turkey Holstein n = 105 Brown Swiss n = 126 n = 63 n = 42 n = 63 n = 63	Early: Day 28–40 Parity: 1 Parity: 2 Parity: 3 Parity: 4	(4.10) (3.13) (3.81) (3.08) (3.38) (3.83)	–	–	–	–	–	HPTLC	Polar lipids ↑ in Holstein compared to Brown Swiss cows. Effect of parity is breed dependent.
X. Zou et al. 2015	Unknown breed Danish n = 10	Colostrum: Day 1–5 MM: Day >16	4.78 5.21	1.20 (25.20) 1.22 (23.49)	1.45 (30.23) 1.92 (36.72)	0.47 (9.89) 0.72 (13.85)	0.35 (7.32) 0.51 (9.83)	1.31 (27.36) 0.84 (16.11)	HPLC, ESLD
de Veth et al. 2016	Holstein Canada n = 5	MM: Day 181–231 Control: RPC12.5: RPC25: A12.5 A25:	–	53.35 51.25 52.98 50.32 52.78	–	–	–	2.91 2.78 2.88 2.63 2.69	HILIC LC-MS
Yao et al. 2016	Holstein China n = 10	MM: Day 30–60		(33.12)	(23.42)	(8.97)	(9.07)	(25.40)	HPLC-ELSD
Z. Liu et al. 2017	Holstein-Friesian Australia n = 19	MM	–	–	–	–	–	–	LC-MS	Highest polar lipid concentration in autumn (April/May) with gradual ↑ in milking season while maintaining a constant proportion of each polar lipid class.
Ali et al. 2018	Egyptian cattle Egypt	nm	–	(48)	(18)	(4)	(12)	(17)	LC-MS
Micek et al. 2019	Holstein-Friesian Poland n = 40 10 cows per treatment group	MM: Day 61–73	EFA-0: 31.3 mg/100 mL milk EFA-1: 24.4 EFA-2: 22.7 EFA-3: 25.0	(43.1) (45.4) (46.4) (46.1)	(33.4) (34.2) (29.9) (33.0)	(10.3) (7.7) (11.5) (7.9)	(8.1) (8.6) (8.2) (8.5)	(5.1) (4.1) (4.0) (4.5)	HPLC
M. Li, Li, et al. 2020	Holstein China n = 30	Colostrum: Day 1–5 MM: 2–3 months		0.129% of total lipids 0.182%	0.261% of total lipids 0.474%	0.031% of total lipids 0.035%	0.035% of total lipids 0.035%	0.608% of total lipids 0.856%	UHPLC-MS

MM: mature milk; nm: not mentioned; TM: transitional milk

^aTotal polar lipid content refers to the cumulation of PC, PE, PI, PS and SM only (5 major polar lipid classes)

^bValue calculated/interpreted based on data provided by author

^cValue represents an average of the dataset

Table 4. Mean ganglioside content (mg/L milk) and proportion of individual gangliosides (mg and % of total gangliosides) in human and bovine milk.

Author	Region	Lactation stage	GM3 mg (or % of total gangliosides)	GD3 mg (or % of total gangliosides)	Total ganglioside mg (or mg/g of fat)	Analysis	Dominant ganglioside	Outcome trend
Human
Full-term
Thakkar et al. 2013	Singapore Female n = 25 Male n = 25	MM: Day 30 Female Male Combined 60 days Female Male Combined 120 days Female Male Combined Average	2.5 2.2 2.3 2.7 3.0 2.9 3.5 4.3 3.9 3.0^b	2.3 2.3 2.3 2.0 1.7 1.9 1.3 2.0 1.7 2.0^b		LC-MS	GM3 GD3 GM3 = GD3 GM3 GM3 GM3 GM3 GM3 GM3	No difference between genders for GM3 & GD3 at 30 days. Greatest ↑ at 120 d which coincides with the largest ↑ in lipid content across the measured timepoints.
Giuffrida et al. 2014	China n = 450	Colostrum/ TM: Day 0–11 MM: 30 days 60 days 120 days	4.3 (53.0^a) 7.4 (81.3^a) 9.1 (91.0^a) 9.8 (91.6^a)	3.8 (47.0^a) 1.7 (18.7^a) 0.9 (9.0^a) 0.9 (8.4^a)	8.1 9.1 10.0 10.7	LC-ESI-HR-MS	GM3 GM3 GM3 GM3	GD3 and GM3 were comparable at 0–11 days. ↑ GM3 to GD3 ratio over lactation.
Ma, Liu, et al. 2015	China n = 14 n = 20 n = 20 n = 20 n = 20 n = 20	MM: 1 month 2 months 3 months 4 months 6 months 8 months	8.5 (65) 11.3 (62) 17.4 (83) 18.3 (92) 21.4 (93) 18.9 (90)	4.6 (35) 7.0 (38) 3.5 (17) 1.5 (8) 1.5 (7) 2.0 (10)	13.1 (0.42) 18.2 (0.49) 20.9 (0.60) 19.8 (0.63) 22.9 (0.64) 21.1 (0.65)	HPLC-MS/MS	GM3 GM3 GM3 GM3 GM3 GM3
Ma, MacGibbon, et al. 2015	Malaysia n = 24 n = 12 n = 48 n = 42 n = 42	Colostrum TM MM: 2 months 6 months 12 months	6.5 (24^a) 8.3 (44^a) 8.3 (56^a) 21.4 (85^a) 13.8 (83^a)	20.3 (76^a) 10.6 (56^a) 6.5 (44^a) 4.3 (17^a) 2.8 (17^a)	26.8 18.9 14.8 25.3 16.6	LC-MS	GD3 GD3 GM3 GM3 GM3
Giuffrida et al. 2016	China n = 539 (total) n = 38 n = 45 n = 30 n = 113 n = 22 n = 21 n = 38 n = 81 n = 120 n = 113 n = 112 n = 345	Colostrum: Day 0–5 Guangzhou Beijing Suzhou Average TM: Day 6–15 Guangzhou Beijing Suzhou Average MM: 0.5–8 months Guangzhou Beijing Suzhou Average	4.0 (60.6) 3.7 (62.7) 4.0 (31.7) 3.8 (47.5) 7.7 (72.0) 3.3 (52.4) 5.4 (65.1) 5.5 (64.7) 10.5 (91.3) 9.0 (92.8) 10.8 (90.8) 10.1 (91.8)	2.8 (42.4) 2.2 (37.3) 8.6 (68.3) 4.1 (51.2) 3.0 (28.0) 3.0 (47.6) 2.9 (34.9) 3.0 (35.3) 1.0 (8.7) 0.7 (7.2) 1.1 (9.2) 1.0 (9.1)	6.6 5.9 12.6 8.0 10.7 6.3 8.3 8.5 11.5 9.7 11.9 11.0	LC-MS	GM3 GM3 GD3 GD3 GM3 GM3 GM3 GM3 GM3 GM3 GM3 GM3	On average, GD3 ↓ and GM3 ↑ over lactation.
McJarrow et al. 2019	United Arab Emirates n = 41 n = 40	TM: Day 5–15 MM: 6 months	9.47 (45) 18.62 (92)	11.71 (55) 1.57 (8)	21.18 20.18	LC-MS	GD3 GM3	GD3 ↓ and GM3 ↑ over lactation.
Reported range^b		MM	2.2–21.4	0.7–20.3	5.9–26.8
Bovine
Sørensen 2006	Danish Holstein raw milk (n = 7) Jersey raw milk (n = 7) Pasteurised skimmed milk from retail storages (n = 4)	nm	0.10 (1.1^a) 0.18 (1.0^a) 0.07 (1.4^a)	9.3 (98.9^a) 17.5 (98.9^a) 5.0 (98.6^a)	9.4 17.7 5.1	LC-MS	GD3 GD3 GD3
Lee et al. 2013	USA Holstein n = 1	Colostrum: Day 2 TM: Day 15 MM: Day 90	0.98 (6.1^a) 0.15 (4.3^a) 0.15 (5.9^a)	15.2 (93.9^a) 3.3 (95.7^a) 2.4 (94.1^a)	16.18^b 3.45^b 2.55^b	UHPLC-MS	GD3	More gangliosides in colostrum, which dramatically ↓ over lactation.
Rivas-Serna et al. 2015	Canada Holstein	Colostrum Whole milk	0.35 0.01	4.54 3.17	6.64 mg/L 3.85 mg/L	LC-MS	GD3 GD3

MM: mature milk; TM: transitional milk

^aValue calculated/interpreted based on data provided by author

^bAssuming all studies identified in this systematic review reporting LC- MS data and not corrected for possible differences in methodology (predominantly difference in GD3 species detection).

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