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. 2021 Jul 7;12:672532. doi: 10.3389/fendo.2021.672532

Table 4.

Effects of glyphosate (active ingredient) (Gly) and glyphosate-based herbicides (GBHs) associated with female fertility and those reported in the successive generations (F1, F2 or F3) of mammalian offspring after maternal exposure.

Compound tested Exposure type Fertility associated effects F1, F2 or F3 offspring effects Reference
-GBH: Magnum Super II
-Gly
Model: pregnant female rat
Exposure: trough food from gestational day (GD)9 to lactational day (LD)21
Dose: 2 mg Gly/kg bw/day
Target of study: uterus
F1 female rats
GD5:
Gly and GBH:
-increased E2 serum levels
-altered expression of implantation related- molecules: (ERα, Hoxa10 and Lif)
GD19:
Gly and GBH:
-increased rate of preimplantation losses
Gly:
-decreased weight of F1 male pups at birth
(31)
-GBH: Roundup Full II Model: pregnant female rat
Exposure: subcutaneous injections from postnatal day (PND)1 to PND7
Dose: 2 mg Gly/kg bw/day
Target of study: uterus
GD9:
-altered endometrial decidualization process
-increased proliferation (Ki67)
GD19:
-increased rate of postimplantation losses
NR (34)
-GBH: Roundup
-Gly
Model: pregnant female mouse
Exposure: trough water from GD1 to GD19
Concentration: 5 mg Gly/ml
Target of study: ovary, hypothalamus and pituitary gland.
GD19:
Gly and GBH:
-decreased ovary weight and altered ovarian histology.
-decreased progesterone serum levels
-altered expression of genes involved in hormonal balance in hypothalamus, pituitary gland and ovary: GnRH, LHR and3β-HSD
-altered serum and ovarian markers of oxidative response
Gly:
-increased estradiol serum levels
Gly or GBH
-specific treatment-related differences
Gly:
-altered sex ratio of F1 fetuses (increased male to female ratio)
(36)
-GBH: Magnum Super II Model: pregnant female rat
Exposure: through food from GD9 to LD21.
Dose: 350 mg Gly/kg bw/day
Target of study: uterus
F1 female rats
GD5:
-increased ERα mRNA expression by increasing ERα-O transcript variant
-Epigenetic alterations in the O promoter of ERα
NR (37)
-Gly Model: pregnant female rat (F0)
Exposure: trough daily intraperitoneal (ip) injections from GD8 to GD14
Dose: 25 mg Gly/kg bw/day
Target of study: multiple organs
-no changes in fertility rates -decreased body weight at weaning in F1 offspring
-increased age at puberty for F1 and F2 generations
-increased obesity in F2 and F3 generations
-increased frequency of histological alterations in F1, F2 and F3 generations (ovary, kidney, testis, prostate)
-parturition abnormalities in F2 and F3 dams
-increased tumor development in F2 generation (mammary adenomas)
-differential methylation genes in sperm from F1, F2 and F3 generations
(38)
-GBH: Magnum Super II Model: pregnant female rat (F0)
Exposure: through food from GD9 to LD21.
Dose: 2 and 200 mg Gly/kg bw/day
Target of study: uterus, placenta, fetal parameters
F1 female rats
GD19:
200 mg Gly/kg bw/day
-increased rate of preimplantation losses
F2 offspring
GD19:
200 mg Gly/kg bw/day:
-decreased fetal weight and length
-increased placental index
-structural congenital anomalies
2 mg Gly/kg bw/day:
-decreased fetal weight
(121, 122)
-GBH: Roundup Model: pregnant female rat
Exposure: by gavage from GD6 to GD15
Dose: 500, 750 and 1000 mg Gly/kg bw/day
Target of study: multiple organs
-no changes associated with female fertility All doses:
-increased occurrence of delayed ossification in several body structures in F1 fetuses
(123)
-GBH: Roundup Full II
-Mixture (MIX): commercial formulations of Gly + endosulfan (Endo) 
Model: neonatal and pregnant female rat
Exposure: subcutaneous injections from PND1 to PND7
Dose: 2 mg Gly/kg bw/day or 2 mg Gly/kg bw/day + 600 µg Endo/kg bw/day
Target of study: uterus
PND8 :
GBH and MIX:
-increased incidence of luminal epithelial hyperplasia
-increased PR and Hoxa10 protein expression
GD19:
GBH and MIX:
-increased rate of postimplantation losses.
NR (127)
-GBH: Roundup
-Mixture (MIX): commercial formulations of Gly + Paraquat (Pq)
Model: pregnant female rat
Exposure: subcutaneous injections from GD1 to GD7
Dose: 500 mg Gly/kg bw/day or 500 mg Gly/kg bw/day + 50 mg Pq/kg bw/day
Target of study: uterus, ovary and embryonic cells
GD7:
GBH and MIX:
-decreased body and ovary weight
-decreased number of corpora lutea
-increased rate of preimplantation losses
-decreased surface and glandular epithelia and diameter of endometrial glands
MIX:
-disorganization of the cytotrophoblast and cell degeneration within the blastocyte cavity
NR (128)
-GBH: Kalach 360 SL Model: adult female rat
Exposure: trough water during 60 days
Dose: 126 and 315 mg Gly/kg bw/day
Target of study: ovary
At the end of the treatment:
-impaired folliculogenesis and ovary development.
-necrosis, vacuolisation of follicles, dissociated oocytes and granulosa cells.
-increased atretic follicles.
-decreased E2 serum levels
-decreased antioxidant enzyme activities: catalase (CAT), superoxide dismutase and glutathione peroxidase (GP).
NR (129)
-GBH: Roundup Bioflow
-Gly
Model: pregnant female rat
Exposure: through drinking water from GD6 to 6-weeks or 13 weeks after weaning
Dose: 1.75 mg Gly/kg bw/day
Target of study: multiple targets
NR F1 offspring
Gly and GBH:
-altered developmental parameters (anogenital distance, age at first estrous)
-altered sexual hormone concentrations
-specific sex-related and treatment-related differences
(173)
-GBH: Roundup-Gly Model: pregnant female mouse
Exposure: trough drinking water from GD1 to GD21
Dose: 5 mg Gly/ml
Target of study: liver
NR F1 offspring
Gly and GBH:
-decreased body weight at weaning
-disruption of lipid metabolism
(174)
-GBH: Glifloglex Model: pregnant female rat
Exposure: trough drinking water from GD1 to LD21
Dose: 100 and 200 mg Gly/kg bw/day
Target of study: brain
NR F1 offspring
-early onset of cliff aversion reflex and auditory canal opening.
-decreased locomotor activity and anxiety levels
-exacerbated emotionality
(175)
-GBH: Glifloglex Model: pregnant female rat
Exposure: trough drinking water from GD1 to LD21
Dose: 100 and 200 mg Gly/kg bw/day
Target of study: brain
NR F1 offspring
-altered antioxidant status
-altered enzymes activity involved in glutamatergic and cholinergic systems
-impairment in recognition memory
(176)
-GBH: Roundup Model: pregnant female rat
Exposure: trough oral gavage from GD0 to LD21
Dose: 250 and 500 mg Gly/kg bw/day
Target of study: brain
-decreased fertility rate and gestational index F1 offspring
-decreased number of pups per litter
-decreased body weight on PND15 and PND21
Offspring: delayed reflexes and altered motor development
Adults: decreased locomotor activity, sociability, learning and impaired memory
(177)
-Gly Model: pregnant female rat
Exposure: ip injections every 48 hs from GD8 to GD20
Dose: 24 and 35 mg Gly/kg bw/day
Target of study: brain
NR F1 offspring
-decreased body weight in pups from PND19
-altered dose-dependent reflexes development, motor activity and cognitive function
-altered Wnt5a-CaMKII pathway in fetal hippocampus
(178)
-GBH: Roundup Transorb Model: pregnant female rat
Exposure: by oral gavage from GD18 to LD5
Dose: 5 and 50 mg Gly/kg bw/day
Target of study: brain (cerebellum and cortex)
NR F1 male offspring (PND90)
-altered gene expression involved in oxidant defense, inflammation and lipid metabolism
-correlation of changes in gene expression with serum concentrations of oxidative stress-related metabolites
(179)
-GBH: Roundup Maxload Model: pregnant female mouse
Exposure: trough drinking water from GD5 to LD21
Dose: 12.5 mg Gly/kg bw/day
Target of study: brain (prefrontal cortex, hippocampus and striatum)
NR F1 juvenile offspring (PND28-PND35)
-autism spectrum disorder (ASD)-like behavioral abnormalities (cognitive and social interaction deficits)
-increased soluble epoxide hydrolase (sEH) expression
-ASD-like behaviors prevented by oral administration of an sEH inhibitor
-abnormal composition of gut microbiota
(180)

3β-HSD, 3β-hydroxysteroid dehydrogenase; ASD, autism spectrum disorder; CaMKII, Ca2+/calmodulin-dependent protein kinase; CAT, catalase; Endo, endosulfan; ERα, estrogen receptor alpha; GBH, glyphosate based-herbicide; GD, gestational day; Gly, glyphosate (active ingredient); GP, glutathione peroxidase; Hoxa10, Homeobox A10; ip, intraperitoneal; LD, lactational day; Lif, Leukemia inhibitory factor; LHR, luteinizing hormone receptor; MIX, mixture; NR, Not reported; PND, postnatal day; PR, progesterone receptor; Pq, paraquat; sEH, soluble epoxide hydrolase; Wnt5a, wingless-type MMTV integration site.