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. 2019 Apr 15;14(4):e0215390. doi: 10.1371/journal.pone.0215390

Table 1. Steroid content of testis, prostate, and liver from puberal and postpuberal littermate boars treated with the aromatase inhibitor, letrozole, or with the canola oil vehicle1.

Testis2
C18 V-16 L-16 V-20 L-20 V-40 L-40
Estrone 22.07 (9.92) 1.96 (0.78) 17.03 (8.97) 1.88 (0.49) 9.96 (2.71) 4.48 (2.28)
Estrone-3-SO4 76.11 (29.56) 0.92 (0.45) 50.75 (35.24) 2.02 (2.44) 49.52 (19.75) 46.42 (35.49)
Estrone-3-GLU ND ND ND ND ND ND
16α-Hydroxyestrone 1.70 (0.68) 0.04 (0.04) 0.68 (0.08) 0.13 (0.11) 0.51 (0.31) 0.33 (0.09)
Estradiol 44.45 (29.11) 7.88 (3.74) 56.26 (27.04) 7.86 (4.62) 39.59 (4.83) 19.68 (10.45)
Estradiol-3-SO4 586.5 (246.2) 2.69 (1.14) 381.6 (271.3) 8.30 (8.91) 273.9 (70.6) 199.7 (129.9)
Estradiol-3-GLU 0.27 (0.26) 0.12 (0.18) 0.31 (0.62) 0.03 (0.06) 0.02 (0.04) 0.02 (0.03)
Estradiol-17-GLU 3.64 (3.30) 0.66 (0.77) 2.94 (2.18) 0.97 (1.08) 2.13 (2.13) 0.24 (0.38)
Estriol 0.20 (0.29) trace 0.29 (0.41) 0.01 (0.02) 0.04 (0.08) 0.04 (0.08)
C19 V-16 L-16 V-20 L-20 V-40 L-40
Dehydroepiandrosterone 85.96 (23.65) 132.90 (138.80) 98.91 (105.92) 48.55 (45.99) 52.26 (30.66) 37.13 (32.51)
Dehydroepiandrosterone-SO4 237.8 (43.87) 163.7 (179.6) 193.7 (71.79) 125.9 (70.36) 98.37 (15.06) 102.0 (55.65)
5-Androstenediol 75.40 (35.83) 104.3 (107.7) 86.29 (113.7) 37.4 (30.84) 84.27 (88.43) 28.20 (22.01)
4-Androstenediol 5.58 (4.64) 5.01 (4.81) 5.35 (6.13) 1.89 (1.21) 4.77 (3.41) 2.20 (1.87)
Androstenedione 58.11 (31.33) 36.20 (32.91) 53.49 (32.36) 21.24 (9.45) 18.20 (12.55) 13.08 (4.79)
19-Hydroxyandrostendione 12.80 (14.66) 0.35 (0.34) 10.60 (11.26) 0.93 (0.99) 2.61 (1.42) 3.07 (2.78)
Testosterone 236.2 (130.6) 157.8 (192.4) 166.5 (92.85) 84.42 (59.00) 61.86 (36.59) 36.84 (12.43)
Epitestosterone 4.50 (2.95) 2.42 (3.65) 4.81 (2.94) 2.32 (0.76) 1.01 (0.74) 0.67 (0.49)
Epi/Testosterone-SO4 12.60 (6.59) 11.32 (4.70) 8.44 (6.36) 4.27 (2.42) 4.56 (3.92) 7.53 (6.41)
5α-Dihydrotestosterone 1.87 (1.88) 4.39 (6.76) 2.42 (2.58) 3.12 (3.31) 0.54 (0.63) 1.33 (1.09)
6-Ketotestosterone 59.83 (96.59) 55.22 (75.38) 45.08 (55.70) 33.28 (27.30) 7.44 (11.89) 5.88 (5.80)
6-Dehydrotestosterone 0.86 (0.46) 0.40 (0.25) 0.29 (0.15) 0.30 (0.09) 0.21 (0.12) 0.22 (0.13)
17β-Dihydroepiandrosterone 181.7 (162.6) 174.8 (242.7) 151.0 (170.3) 98.98 (46.79) 130.6 (93.27) 82.13 (51.50)
6α-Hydroxytestosterone 12.16 (8.52) 3.54 (6.53) 9.13 (6.32) 2.02 (3.49) 1.49 (0.90) 0.88 (0.79)
Adrenosterone 2.74 (2.82) 2.37 (2.47) 1.40 (0.89) 1.49 (1.15) 0.39 (0.45) 0.40 (0.29)
Epiandrosterone 12.85 (7.10) 15.66 (26.94) 10.91 (7.72) 10.13 (14.69) 3.81 (4.12) 6.51 (5.76)
C21
Pregnenolone 835.4 (357.8) 822.5 (462.5) 739.8 (594.0) 806.3 (725.1) 484.4 (107.0) 397.8 (239.8)
Pregnenolone-3-SO4 59.3 (11.5) 130.6 (177.8) 66.7 (52.8) 87.7 (77.3) 39.8 (20.9) 38.5 (33.6)
17-Hydroxypregnenolone 396.0 (265.0) 406.5 (360.5) 421.4 (507.5) 395.9 (536.0) 255.1 (208.4) 214.1 (147.9)
Progesterone 6.92 (2.26) 10.30 (6.50) 9.03 (9.34) 16.86 (23.74) 5.08 (4.11) 4.14 (2.98)
17-Hydroxyprogesterone 2.52 (1.01) 4.04 (2.70) 3.48 (3.44) 4.68 (4.61) 2.89 (2.13) 2.08 (1.08)
11-Deoxycortisol 2.88 (2.15) 1.07 (0.72) 2.25 (1.42) 0.95 (0.71) 0.97 (0.55) 0.49 (0.30)
Cortisol 4.14 (4.13) 0.69 (0.79) 3.27 (3.09) 1.01 (0.99) 2.10 (1.49) 2.09 (1.39)
Deoxycorticosterone 1.59 (1.22) 0.50 (0.43) 1.35 (1.02) 0.44 (0.26) 0.44 (0.35) 0.27 (0.11)
Corticosterone 0.27 (0.52) 0.41 (0.43) 1.02 (1.81) 0.16 (0.12) 0.37 (0.27) 0.42 (0.27)
Epiallopregnanolone 78.13 (62.52) 120.8 (147.3) 81.82 (97.05) 162.6 (179.2) 81.87 (95.14) 47.55 (25.79)
Allo/Pregnanolone-SO4 1.59 (1.36) 3.78 (3.38) 0.93 (0.61) 1.90 (1.84) 0.42 (0.33) 0.23 (0.18)
Epi/Epiallopregnanolone-SO4 2.91 (1.80) 21.39 (35.92) 7.94 (11.73) 14.97 (18.26) 4.37 (5.82) 3.15 (3.72)
Prostate3
C18 V-16 L-16 V-20 L-20 V-40 L-40
Estrone 3.64 (1.86) 1.59 (0.51) 2.14 (0.92) 2.04 (1.18) 2.94 (2.15) 1.65 (0.49)
Estrone-3-SO4 9.37 (7.78) 0.36 (0.19) 5.27 (2.42) 0.49 (0.12) 9.32 (8.19) 4.00 (3.01)
Estrone-3-GLU ND ND ND ND ND ND
16α-Hydroxyestrone ND ND ND ND ND ND
Estradiol 18.33 (8.28) 11.80 (5.35) 15.50 (10.56) 13.34 (7.89) 19.43 (14.06) 8.04 (1.98)
Estradiol-3-SO4 7.44 (5.38) 0.54 (0.42) 4.28 (1.64) 0.73 (0.99) 10.86 (13.35) 3.25 (1.92)
Estradiol-3-GLU 0.35 (0.57) 0.002 (0.002) 0.05 (0.09) 0.12 (0.17) 1.31 (0.99) 0.02 (0.03)
Estradiol-17-GLU 0.42 (0.72) 0.002 (0.001) 0.20 (0.40) 1.51 (2.00) 1.40 (1.23) 0.94 (0.86)
Estriol ND ND ND ND ND ND
C19
Dehydroepiandrosterone ND ND ND ND ND ND
Dehydroepiandrosterone-SO4 29.29 (19.57) 9.89 (7.27) 15.36 (5.18) 12.16 (7.53) 31.03 (24.15) 13.15 (10.51)
5-Androstenediol ND ND ND ND ND ND
4-Androstenediol ND ND ND ND ND ND
Androstenedione 0.26 (0.10) 0.24 (0.09) 0.22 (0.10) 0.13 (0.06) 0.38 (0.29) 0.13 (0.01)
19-Hydroxyandrostendione ND ND ND ND ND ND
Testosterone 1.67 (0.32) 1.04 (1.67) 0.36 (0.15) 0.55 (0.29) 1.07 (0.76) 0.33 (0.15)
Epitestosterone 0.01 (0.02) 0.09 (0.07) 0.02 (0.02) 0.09 (0.10) 0.27 (0.53) 0.27 (0.28)
Epi/Testosterone-SO4 ND ND ND ND ND ND
5α-Dihydrotestosterone 2.51 (1.10) 1.79 (1.17) 1.29 (0.84) 1.23 (0.39) 0.93 (0.95) 0.99 (0.49)
6-Ketotestosterone ND ND ND ND ND ND
6-Dehydrotestosterone ND ND ND ND ND ND
17β-Dihydroepiandrosterone trace trace trace trace trace trace
6α-Hydroxytestosterone ND ND ND ND ND ND
Adrenosterone ND ND ND ND ND ND
Epiandrosterone 4.76 (2.82) 6.24 (5.94) 6.01 (4.52) 7.01 (4.62) 4.41 (3.25) 3.27 (2.01)
C21
Pregnenolone ND ND ND ND ND ND
Pregnenolone-3-SO4 0.49 (0.39) 0.36 (0.29) 0.42 (0.19) 1.29 (0.88) 0.98 (0.80) 0.62 (0.55)
17-Hydroxypregnenolone ND ND ND ND ND ND
Progesterone 0.70 (0.34) 0.31 (0.21) 0.73 (0.86) 1.64 (2.36) 0.58 (0.50) 0.25 (0.20)
17-Hydroxyprogesterone ND ND ND ND ND ND
11-Deoxycortisol ND ND ND ND ND ND
Cortisol 2.85 (1.87) 0.86 (0.72) 3.13 (4.05) 3.39 (3.70) 3.38 (3.57) 1.12 (0.69)
Deoxycorticosterone 0.24 (0.36) 0.11 (0.10) 0.24 (0.26) 0.24 (0.25) 0.07 (0.11) 0.08 (0.07)
Corticosterone 1.84 (1.38) 0.36 (0.50) 0.82 (1.15) 2.32 (3.12) 0.26 (0.17) 0.35 (0.28)
Epiallopregnanolone ND ND ND ND ND ND
Allo/Pregnanolone-SO4 ND ND ND ND ND ND
Epi/Epiallopregnanolone-SO4 trace trace trace trace trace trace
Liver4
C18 V-16 L-16 V-20 L-20 V-40 L-40
Estrone 0.22 (0.03) 0.33 (0.13) 0.53 (0.20) 0.41 (0.05) 0.48 (0.21) 0.29 (0.12)
Estrone-3-SO4 4.84 (3.40) 0.06 (0.04) 6.02 (1.51) 0.26 (0.39) 6.85 (4.79) 2.18 (1.30)
Estrone-3-GLU 1.06 (0.36) 0.12 (0.22) 0.14 (0.13) 0.36 (0.44) 1.40 (2.23) 0.90 (1.01)
16α-Hydroxyestrone 0.01 (0.003) 0.03 (0.01) 0.02 (0.01) 0.02 (0.01) 0.02 (0.01) 0.02 (0.004)
Estradiol 1.27 (0.05) 1.68 (0.84) 2.89 (2.07) 1.47 (0.50) 2.40 (0.80) 1.43 (1.06)
Estradiol-3-SO4 14.26 (15.44) 0.13 (0.08) 34.31 (12.07) 1.35 (1.70) 67.40 (43.92) 21.61 (14.41)
Estradiol-3-GLU 1.19 (1.33) 0.004 (0.01) 0.95 (0.88) 0.02 (0.02) 3.05 (3.03) 2.18 (1.72)
Estradiol-17-GLU 1.11 (1.57) 0.32 (0.64) 0.57 (0.47) 0.13 (0.15) 1.79 (0.29) 0.62 (0.67)
Estriol 0.07 (0.10) trace 0.16 (0.23) trace 0.01 (0.02) 0.21 (0.34)
C19 V-16 L-16 V-20 L-20 V-40 L-40
Dehydroepiandrosterone ND ND ND ND ND ND
Dehydroepiandrosterone-SO4 11.14 (4.38) 6.96 (3.49) 16.70 (4.33) 11.66 (10.59) 17.78 (14.61) 9.23 (6.83)
5-Androstenediol ND ND ND ND ND ND
4-Androstenediol ND ND ND ND ND ND
Androstenedione 0.07 (0.01) 0.13 (0.05) 0.34 (0.23) 0.17 (0.08) 0.11 (0.09) 0.07 (0.04)
19-Hydroxyandrostendione ND ND ND ND ND ND
Testosterone 0.19 (0.01) 0.31 (0.27) 0.43 (0.05) 0.65 (0.32) 0.60 (0.49) 0.31 (0.08)
Epitestosterone 0.26 (0.08) 0.18 (0.15) 0.14 (0.11) 0.25 (0.23) 0.20 (0.21) 0.19 (0.19)
Epi/Testosterone-SO4 2.35 (2.00) 2.15 (2.10) 2.87 (0.76) 2.90 (3.31) 4.93 (2.03) 4.79 (4.31)
5α-Dihydrotestosterone 0.04 (0.06) 0.09 (0.10) 0.04 (0.07) 0.01 (0.02) 0.13 (0.15) 0.05 (0.07)
6-Ketotestosterone ND ND ND ND ND ND
6-Dehydrotestosterone ND ND ND ND ND ND
17β-Dihydroepiandrosterone ND ND ND ND ND ND
6α-Hydroxytestosterone ND ND ND ND ND ND
Adrenosterone ND ND ND ND ND ND
Epiandrosterone ND ND ND ND ND ND
C21 V-16 L-16 V-20 L-20 V-40 L-40
Pregnenolone ND ND ND ND ND ND
Pregnenolone-3-SO4 0.60 (0.02) 0.71 (0.86) 0.87 (0.54) 1.07 (0.64) 0.95 (0.39) 0.96 (0.76)
17-Hydroxypregnenolone ND ND ND ND ND ND
Progesterone 0.06 (0.03) 0.17 (0.11) 1.09 (1.16) 1.05 (1.09) 0.30 (0.06) 0.30 (0.14)
17-Hydroxyprogesterone ND ND ND ND ND ND
11-Deoxycortisol 0.14 (0.08) 0.28 (0.25) 2.04 (2.09) 1.87 (1.86) 0.36 (0.27) 0.30 (0.19)
Cortisol 0.47 (0.06) 1.64 (2.25) 13.23 (12.51) 7.73 (8.98) 1.40 (1.14) 1.70 (1.05)
Deoxycorticosterone 0.03 (0.01) 0.04 (0.04) 0.38 (0.33) 0.36 (0.45) 0.11 (0.08) 0.07 (0.05)
Corticosterone 0.20 (0.21) 0.30 (0.26) 0.60 (0.54) 0.86 (0.91) 0.07 (0.06) 0.12 (0.11)
Epiallopregnanolone ND ND ND ND ND ND
Allo/Pregnanolone-SO4 ND ND ND ND ND ND
Epi/Epiallopregnanolone-SO4 trace trace trace trace trace trace

1Values represent means (SD) in ng/g. Values in bold indicate statistically significant differences between letrozole-treated boars and vehicle-treated littermates at the indicated age.

2Values for testis tissue at each age are based upon one boar from each of four litters treated with letrozole from 11 to 16 weeks of age and littermates treated with the canola oil vehicle. All boars came from a total of five litters.

3Values for prostate tissue at each age are based upon one boar from each of four litters treated with letrozole from 11 to 16 weeks of age (three boars treated with vehicle at 16 weeks of age and three boars treated with letrozole at 20 weeks of age) and littermates treated with the canola oil vehicle. Tissues came from same boars that donated testicular tissue.

4Values for liver tissue at each age are based upon one boar from each of four litters treated with letrozole from 11 to 16 weeks of age (three boars treated with vehicle at 16 weeks of age and at 20 weeks of age) and littermates treated with the canola oil vehicle. Tissues came from same boars that donated testicular tissue.