FIPAT TE2 Part1
FIPAT TE2 Part1
FIPAT TE2 Part1
Second Edition
International Embryological Terminology
FIPAT
The Federative International Programme for Anatomical Terminology
A programme of the International Federation of Associations of Anatomists (IFAA)
TE2, PART I
Contents
Published pending approval by the General Assembly at the next Congress of IFAA (2019)
The individual terms in this terminology are within the public domain. Statements about terms being part of this international standard terminology should use
the above bibliographic reference to cite this terminology. The unaltered PDF files of this terminology may be freely copied and distributed by users. IFAA
member societies are authorized to publish translations of this terminology. Authors of other works that might be considered derivative should write to the
Chair of FIPAT for permission to publish a derivative work.
Caput I: NOMINA GENERALIA Chapter 1: GENERAL TERMS
160 Mensurae embryonicae et Embryonic and fetal Embryonic and fetal Endnote 12
fetales measurements measurements
161 Aetas a fecundatione Fertilization age Fertilization age Endnote 13
162 Aetas ab ovulatione Ovulation age Ovulation age Endnote 14
163 Aetas ab inseminatione Insemination age Insemination age Endnote 15
164 Hebdomades post coitum Coital weeks Coital weeks Endnote 16
165 Hebdomades post menses ultimas Menstrual weeks Menstrual weeks Endnote 17
166 Longitudo corona calx Crown-heel length Crownheel length CHL; Total length; Standing height
167 Longitudo maxima Greatest length Greatest length GL Endnote 18
168 Longitudo corona nates Crown-rump length Crownrump length CRL; Sitting height
169 Longitudo cervix nates Neck-rump length Neckrump length
170 Longitudo ossis femoris ossificati Ossified femur length Ossified femur length
171 Longitudo pedis Foot length Foot length FL
172 Diameter biparietalis Biparietal diameter Biparietal diameter
173 Diameter cavitatis amnioticae Amniotic cavity diameter Amniotic cavity diameter
174 Diameter cavitatis chorionicae Chorionic cavity diameter Chorionic cavity diameter
175 Diameter vesiculae umbilicalis Diameter sacci vitellini Umbilical vesicle diameter Umbilical vesicle diameter Yolk sac diameter
176 Circumferentia abdominis Abdominal circumference Abdominal circumference
177 Circumferentia capitis Head circumference Head circumference
178 Pondus corporis Body weight Body weight
179 Pondus encephali Brain weight Brain weight
180 Pondus placentae Placental weight Placental weight
319 Folliculus ovaricus primarius Primary ovarian follicle Primary ovarian follicle
320 Folliculus ovaricus secundarius Secondary ovarian follicle Secondary ovarian follicle
337 PELLUCIDAGENESIS Zonagenesis ZONA PELLUCIDA FORMATION ZONA PELLUCIDA FORMATION Endnote 22
338 Epithelium simplex cuboideum Simple cuboidal epithelium of Simple cuboidal epithelium of
folliculi ovarici ovarian follicle ovarian follicle
339 Oocytus primarius Primary oocyte Primary oocyte Diploid (2N, 4C)
340 Zona pellucida Zona pellucida Zona pellucida
341 Processus cellulae epithelialis Process of cuboidal epithelial Process of cuboidal epithelial
follicularis cuboidalis follicular cell follicular cell
342 Processus oocyti Process of oocyte Process of oocyte
343 Proteina zonae pellucidae I - III Zona pellucida proteins 1-3 Zona pellucida proteins 1-3 ZP 1-3
717 TEMPUS EMBRYONICUM Gradus carnegienses [St.1 ad 23] EMBRYONIC PERIOD EMBRYONIC PERIOD Carnegie stages [St. 1-23] Endnote 68
718 Embryo Embryo Embryo
719 Embryo pregastrulationis Pregastrulation embryo Pregastrulation embryo Endnote 69
720 Embryo preimplantationis Pre-implantation embryo Preimplantation embryo
721 Embryo preblastocysticus Preblastocystic embryo Preblastocystic embryo
722 Gradus cellulae unicae Embryo unicellularis One-cell stage One cell stage Single cell embryo
723 Oocytus penetratus Oocytus definitivus; Embryo Penetrated oocyte Penetrated oocyte Definitive oocyte; Primordial
primordialis embryo
724 Zygotum Embryo syngamicus Zygote Zygote Syngamic embryo
725 Zona pellucida Zona pellucida Zona pellucida
726 Spatium subzonale Spatium subcapsulare Subzonal space Subzonal space Subcapsular space Endnote 70
727 Pronucleus Pronucleus Pronucleus
728 Pronucleus femininus Pronucleus maternus Female pronucleus Female pronucleus Maternal pronucleus
729 Corpus polare secundum Polocytus secundarius Second polar body Second polar body Second polocyte
730 Reactio corticalis Cortical reaction Cortical reaction
731 Reactio zonalis Reactio capsularis Zonal reaction Zonal reaction Capsular reaction Endnote 71
732 Degradatio mitochondriorum Degradation of paternal Degradation of paternal Endnote 72
paternalium mitochondria mitochondria
733 Conus fertilisationis Fertilization cone Fertilization cone IVF
734 Fusus anaphasis II Anaphase spindle II Anaphase spindle II IVF
735 Fusus telophasis II Telophase spindle II Telophase spindle II IVF
736 Intercorpus Interbody Interbody IVF
Endnote 73
737 Ootidium Ovum; Embryo pronuclearis Ootid Ootid Ovum; Pronuclear embryo
738 Pronucleus masculinus Pronucleus paternus Male pronucleus Male pronucleus Paternal pronucleus
739 Pronuclei admoti Approximated pronuclei Approximated pronuclei
740 Corpusculum precursorium Nucleolar precursor body Nucleolar precursor body
nucleolorum
741 Polus animalis Polus embryonicus presumptivus Animal pole Animal pole Presumptive embryonic pole
742 Zygotum Zygote Zygote
743 Zygotum findens Cleaving zygote Cleaving zygote Endnote 74
833 Crista previllosa mesoblasti Previllous crest of mesoblast Previllous crest of mesoblast Endnote 97
834 Reticulum extraembryonicum Magma reticulare Extra-embryonic reticulum Extraembryonic reticulum Mesenchymal reticulum Endnote 98
835 Blastocystis invadens cum Invading blastocyst with Invading blastocyst with
lacunis trophoblasticis isolated trophoblastic lacunae isolated trophoblastic lacunae
separatis
836 Lacuna trophoblastica Trophoblastic lacuna Trophoblastic lacuna
837 Aggregatio previllosa Previllous clump of cytotrophoblast Previllous clump of cytotrophoblast
cytotrophoblasti
838 Cavitas amniotica definitiva Definitive amniotic cavity Definitive amniotic cavity
839 Chorion primordiale Primordial chorion Primordial chorion
840 Endoblastus extraembryonicus Membrana exocoelomica Extra-embryonic endoblast Extraembryonic endoblast Exocoelomic membrane; Primary Endnote 99
endoderm
841 Vesicula umbilicalis primaria Saccus vitellinus primarius Primary umbilical vesicle Primary umbilical vesicle Primary yolk sac Endnote 100
842 Cavitas vesiculae umbilicalis Cavitas sacci vitellini primarii Cavity of primary umbilical vesicle Cavity of primary umbilical vesicle Cavity of primary yolk sac Endnote 100
primariae
843 Blastocystis invadens cum Invading blastocyst with Invading blastocyst with
lacunis communicantibus intercommunicating lacunae intercommunicating lacunae
844 Obturamentum fibrosus in loco Obturamentum occludens Fibrous coagulum at implantation Fibrous coagulum at implantation Closing plug
implantationis site site
845 Circulus lacunosus vascularis Lacunar vascular circle Lacunar vascular circle Endnote 101
846 Cavitas chorionica primordialis Primordial chorionic cavity Primordial chorionic cavity
992 Cellulae prestaminales Cellulae proprecursoriae; Cellulae Stem cells in early development Stem cells in early development
pluripotentes
993 Cellula externa morulae Outer cell of morula Outer cell of morula
994 Cellula staminalis trophoblastica Trophoblastic stem cell Trophoblastic stem cell TSC
995 Cellula interna morulae Inner cell of morula Inner cell of morula
996 Cellula staminalis embryonica Stipitoblastus Embryonic stem cell Embryonic stem cell ESC
997 Cellula epiblastica Epiblastic cell Epiblastic cell
998 Cellula hypoblastica Hypoblastic cell Hypoblastic cell
1002 Cellulae staminales Cellulae multipotentes et Lineage-restricted stem cells Lineage restricted stem cells Multipotent and unipotent cells Endnote 138
unipotentes
1003 Cellula staminalis somatica Somatic stem cell Somatic stem cell
1004 Cellula stromalis medullae rubrae Bone marrow stromal cell Bone marrow stromal cell
1005 Cellula staminalis medullae osseae Bone marrow stem cell Bone marrow stem cell
1006 Cellula staminalis sanguinis Cord blood stem cell Cord blood stem cell
funiculi umbilicalis
1007 Cellula staminalis umbilicalis Umbilical stem cell Umbilical stem cell
1008 Cellula staminalis conjunctivalis Conjunctival stem cell Conjunctival stem cell
1009 Cellula staminalis cornealis Corneal stem cell Corneal stem cell
1010 Cellula staminalis endothelialis Endothelial stem cell Endothelial stem cell
1011 Cellula staminalis ependymalis Ependymal stem cell Ependymal stem cell
1012 Cellula staminalis epidermalis Epidermal stem cell Epidermal stem cell
1013 Cellula staminalis gastrointestinalis Gastro-intestinal stem cell Gastrointestinal stem cell
1014 Cellula staminalis haematopoietica Haematopoietic stem cell Hematopoietic stem cell
1015 Cellula staminalis Hepatopancreatic stem cell Hepatopancreatic stem cell
hepatopancreatica
1016 Cellula staminalis hypophysialis Hypophysial stem cell Hypophysial stem cell
1017 Cellula staminalis mesechymatica Mesenchymal stem cell Mesenchymal stem cell hMSC
1018 Cellula staminalis myogenica Myogenic stem cell Myogenic stem cell
1019 Cellula staminalis nervosa Neural stem cell Neural stem cell
1020 Cellula staminalis neuronalis Neuronal stem cell Neuronal stem cell
1021 Cellula staminalis glialis Glial stem cell Glial stem cell
1022 Cellula staminalis epidermalis Epidermal neural crest cell stem Epidermal neural crest cell stem eNCSC
cristae neuralis cell cell
1023 Cellula staminalis olfactoria Olfactory stem cell Olfactory stem cell
1024 Cellula staminalis spermatogonica Spermatogonial stem cell Spermatogonial stem cell
1025 Cellula staminalis germinalis Germinal stem cell Germinal stem cell Endnote 139
1026 Cellulae progenetrices Progenitor cells Progenitor cells See relevant tissue derivatives
1070 COMPLEXUS CRISTAE OPTIC NEURAL CREST OPTIC NEURAL CREST Endnote 145
NEURALIS OPTICAE COMPLEX COMPLEX
1071 Mesenchyma oculi Optic mesenchyme Optic mesenchyme In part
1072 Basicranium anterius Anterior basicranium Anterior basicranium In part
1073 Leptomeninx Leptomeninx Leptomeninx
1074 Tunica fibrosa bulbi Fibrous layer of eyeball Fibrous layer of eyeball
1075 Cellula staminalis cornealis Corneal stem cell Corneal stem cell Endnote 146
1076 Melanocytus Melanocyte Melanocyte
1077 Tunica vasculosa bulbi Uvea Vascular layer of eyeball Vascular layer of eyeball Uvea
1078 Pigmentocytus uvealis Pigment cell of uvea Pigment cell of uvea
1079 CRISTA NEURALIS PREOTICA PRE-OTIC NEURAL CREST PREOTIC NEURAL CREST
1080 Complexus cristae neuralis Mesencephalic neural crest Mesencephalic neural crest Endnote 147
mesencephalicae complex complex
1081 Mesenchyma frontonasale Frontonasal mesenchyme Frontonasal mesenchyme
1082 Basicranium anterius Anterior basicranium Anterior basicranium In part
1083 Os frontale Frontal bone Frontal bone
1084 Pars squamosa ossis temporalis Squamous part of temporal bone Squamous part of temporal bone
1085 Viscerocranium membranaceum Membranous viscerocranium Membranous viscerocranium In part
1086 Leptomeninx Leptomeninx Leptomeninx
1087 Mesenchyma oculi Optic mesenchyme Optic mesenchyme In part
1088 Cellula staminalis cornealis Corneal stem cell Corneal stem cell
1089 Keratocytus Keratocyte Keratocyte
1090 Adipocytus Adipocyte Adipocyte
1091 Melanocytus Melanocyte Melanocyte
1092 Epithelium posterius corneae Endothelium of anterior chamber Endothelium of anterior chamber
1093 Stroma iridis Stroma of iris Stroma of iris
1094 Membrana pupillaris Pupillary membrane Pupillary membrane Iridopupillary membrane In part
1095 Ectomesenchyma dentale Dental ectomesenchyme Dental ectomesenchyme
1096 Papilla dentis Dental papilla Dental papilla
1097 Odontoblastus Odontoblast Odontoblast
1098 Crista neuralis isthmica Isthmic neural crest Isthmic neural crest Endnote 148
1099 Via migrationis dorsolateralis Dorsolateral migration pathway Dorsolateral migration pathway
1100 Mesenchyma capitis * Head mesenchyme * Head mesenchyme * In part
1101 Leptomeninx Leptomeninx Leptomeninx
1118 Complexus cristae neuralis Faciovestibulocochlear neural Faciovestibulocochlear neural Endnote 150
faciovestibulocochlearis crest complex crest complex
1119 Complexus cristae neuralis Facial neural crest complex Facial neural crest complex Endnote 151
facialis
1120 Via migrationis dorsolateralis Dorsolateral migration pathway Dorsolateral migration pathway
1121 Leptomeninx Leptomeninx Leptomeninx
1122 Mesenchyma capitis Head mesenchyme Head mesenchyme In part
1123 Dermis cervicalis Dermis of neck Dermis of neck In part
1124 Adipocytus Adipocyte Adipocyte
1125 Melanocytus Melanocyte Melanocyte
1126 Viscerocranium membranaceum Membranous viscerocranium Membranous viscerocranium In part
1127 Cartilago arcus pharyngei Second pharyngeal arch Second pharyngeal arch §Reichert§
secundi cartilage cartilage
1128 Ossiculae auditus Auditory ossicles Auditory ossicles In part
Endnote 152
1129 Os hyoideum Hyoid bone Hyoid bone In part
1130 Fasciae propriae musculorum Second pharyngeal arch Second pharyngeal arch
arcus pharyngei secundi muscle sheaths muscle sheaths
1131 Via migrationis ventrolateralis Ventrolateral migration pathway Ventrolateral migration pathway
1132 Neuron sensorium ganglii Geniculate ganglion cell Geniculate ganglion cell
geniculi
1133 Gliocytus ganglionicus Ganglionic satellite cell Ganglionic satellite cell
1134 Schwannocytus Schwann cell Schwann cell
1135 COMPLEXUS CRISTAE OTIC NEURAL CREST OTIC NEURAL CREST Endnote 153
NEURALIS OTICAE COMPLEX COMPLEX
1136 Via migrationis dorsolateralis Dorsolateral migration pathway Dorsolateral migration pathway
1137 Leptomeninx Leptomeninx Leptomeninx
1146 CRISTA NEURALIS POSTOTICA POSTOTIC NEURAL CREST POSTOTIC NEURAL CREST
1147 Complexus cristae neuralis Glossopharyngeal neural crest Glossopharyngeal neural crest Endnote 154
glossopharyngealis complex complex
1148 Via migrationis dorsolateralis Dorsolateral migration pathway Dorsolateral migration pathway
1149 Leptomeninx Leptomeninx Leptomeninx
1150 Crista neuralis cardiaca Cardiac neural crest Cardiac neural crest In part
Endnote 155
1151 Cartilago arcus pharyngei tertii Third pharyngeal arch cartilage Third pharyngeal arch cartilage
1152 Os hyoideum Hyoid bone Hyoid bone In part
1153 Fascia propria musculi Stylopharyngeus muscle sheath Stylopharyngeus muscle sheath
stylopharyngei
1154 Via migrationis ventrolateralis Ventrolateral migration pathway Ventrolateral migration pathway
1155 Neuron sensorium ganglii Glossopharyngeal ganglion cell Glossopharyngeal ganglion cell
glossopharyngei
1156 Ganglion glossopharyngeum Superior glossopharyngeal Superior glossopharyngeal
superius ganglion ganglion
1157 Ganglion glossopharyngeum Inferior glossopharyngeal Inferior glossopharyngeal
inferius ganglion ganglion
1158 Gliocytus ganglionicus Ganglionic satellite cell Ganglionic satellite cell
1159 Schwannocytus Schwann cell Schwann cell
1160 Complexus cristae neuralis Vagal neural crest complex Vagal neural crest complex Endnote 154
vagalis
1161 Via migrationis dorsolateralis Dorsolateral migration pathway Dorsolateral migration pathway
1162 Leptomeninx Leptomeninx Leptomeninx
1163 Crista neuralis cardiac Cardiac neural crest Cardiac neural crest In part
1164 Cartilago arcus pharyngei quarti Fourth pharyngeal arch cartilage Fourth pharyngeal arch cartilage
1165 Cartilagines laryngeae Laryngeal cartilages Laryngeal cartilages Endnote 156
1166 Fasciae propriae musculorum Fourth pharyngeal arch muscle Fourth pharyngeal arch muscle
arcus pharyngei quarti sheaths sheaths
1167 Via migrationis ventrolateralis Ventrolateral migration pathway Ventrolateral migration pathway
1168 Neuron sensorium ganglii vagalis Vagal ganglion cell Vagal ganglion cell
1169 Ganglion vagale superius Superior vagal ganglion Superior vagal ganglion
1170 Ganglion vagale inferius Inferior vagal ganglion Inferior vagal ganglion
1171 Gliocytus ganglionicus Ganglionic satellite cell Ganglionic satellite cell
1182 Complexus cristae neuralis Cardiac neural crest complex Cardiac neural crest complex
cardiacus
1183 Arcus pharyngei tertii, quartii et Third, fourth and sixth pharyngeal Third, fourth and sixth pharyngeal
sexti arches arches
1184 Aa. arcuum pharyngeorum Pharyngeal arch arteries Pharyngeal arch arteries Aortic arches Endnote 157
1185 Paraganglion Paraganglion Paraganglion
1186 Paragangliocytus Cellula typi I Paragangliocyte Paragangliocyte Type I cell
1187 Glandulae parathyroideae Parathyroid glands Parathyroid glands In part
1188 Stroma glandulae Parathyroid stroma Parathyroid stroma
parathyroideae
1189 Cor Heart Heart In part
1190 Basis cordis Base of heart Base of heart In part
1191 Ductus tractus communis Common outflow tract of heart Common outflow tract of heart In part
egressionis cordis
1192 Crista endocardiaca septalis Tuber endocardiacum septale Septal ridge Septal ridge Septal cushion; Parietal cushion
1193 Septum aorticopulmonale Aorticopulmonary septum Aorticopulmonary septum In part
1194 Valva aortae Aortic valve Aortic valve Aortic arterial valve In part
1195 Valva trunci pulmonalis Pulmonary valve Pulmonary valve Pulmonary arterial valve In part
1196 Tubera endocardiaca Atrioventricular endocardial Atrioventricular endocardial In part
atrioventricularia cushions cushions
1197 Cardiomyocytus atrialis Endocrine atrial cardiomyocyte Endocrine atrial cardiomyocyte Atrial myo-endocrine cell; Atrial
secretans myoendocrine cell
1198 Trachea et bronchi Trachea and bronchi Trachea and bronchi In part
1199 Neuroendocrinocytus Respiratory neuro-endocrine cell Respiratory neuro-endocrine cell
respiratorius
1200 Gemma thymica Thymic bud Thymic bud In part
1201 Stroma thymi Thymic stroma Thymic stroma
1202 Glandula thyroidea Thyroid gland Thyroid gland In part
1203 Thyrocytus C C thyrocyte C thyrocyte C cell; Parafollicular cell
1204 Crista neuralis nervi accessorii Neural crest of accessory nerve Neural crest of accessory nerve Endnote 158
1208 Crista neuralis hypoglossalis Crista neuralis occipitalis Hypoglossal neural crest Hypoglossal neural crest Occipital neural crest Endnote 159
1209 Via migrationis dorsolateralis Dorsolateral migration pathway Dorsolateral migration pathway
1210 Leptomeninx Leptomeninx Leptomeninx
1211 Adipocytus Adipocyte Adipocyte
1212 Melanocytus Melanocyte Melanocyte
1213 Chorda hypoglossalis Hypoglossal cord Hypoglossal cord In part
1214 Schwannocytus Schwann cell Schwann cell
1215 CRISTA NEURALIS SPINALIS SPINAL NEURAL CREST SPINAL NEURAL CREST Endnote 160
1216 Via migrationis dorsolateralis Dorsolateral migration pathway Dorsolateral migration pathway Endnote 161
1217 Melanocytus Melanocyte Melanocyte
1218 Via migrationis ventrolateralis Ventrolateral migration pathway Ventrolateral migration pathway Endnote 162
1219 Neuron sensorium ganglii spinalis Spinal ganglion cell Spinal ganglion cell
1220 Gliocytus ganglionicus Ganglionic satellite cell Ganglionic satellite cell
1221 Schwannocytus Schwann cell Schwann cell
1222 Via migrationis ventromedialis Ventromedial migration pathway Ventromedial migration pathway Endnote 163
1223 Linea generationis cellularum Sympathosuprarenal cell lineage Sympathosuprarenal cell lineage Sympatho-adrenal cell lineage;
sympathicosuprarenalium Sympathoadrenal cell lineage
1224 Ganglion trunci sympathici Ganglion of sympathetic trunk Ganglion of sympathetic trunk
1225 Gliocytus ganglionicus Ganglionic satellite cell Ganglionic satellite cell
1226 Medulla suprarenalis Suprarenal medulla Suprarenal medulla
1227 Endocrinocytus chromophilus Medullary chromaffin cell Medullary chromaffin cell
medullaris
1228 Ganglion preaorticum Pre-aortic ganglion Preaortic ganglion §Zuckerkandl§
1229 Gliocytus ganglionicus Ganglionic satellite cell Ganglionic satellite cell
1230 Linea generationis cellularum Parasympathetic cell lineage Parasympathetic cell lineage
parasympathicarum
1231 Ganglion parasympathicum Parasympathetic ganglion Parasympathetic ganglion
1232 Gliocytus ganglionicus Ganglionic satellite cell Ganglionic satellite cell
1233 Textus connectivi atque Connective and supporting Connective and supporting
sustinentes tissues tissues
1234 TEXTUS ADIPOSUS ADIPOSE TISSUE ADIPOSE TISSUE
1235 Mesenchyma somiticum Somitic mesenchyme Somitic mesenchyme
1236 Mesenchyma somatopleurale Somatopleuric mesenchyme Somatopleuric mesenchyme Endnote 164
1237 Mesenchyma splanchnopleurale Splanchnopleuric mesenchyme Splanchnopleuric mesenchyme Endnote 165
1238 Ectomesenchyma Mesenchyma cristae neuralis Ectomesenchyme Ectomesenchyme Neural crest mesenchyme
1239 Mesenchyma ex eminentia Mesenchyme from caudal Mesenchyme from caudal
caudale eminence eminence
1240 Cellula adipocytoprogenetrix Fat cell progenitor Fat cell progenitor Endnote 166
1275 Ossificatio membranacea Ossificatio desmalis Membranous ossification Membranous ossification Intramembranous ossification
1276 Mesenchyma blastemale Blastemal mesenchyme Blastemal mesenchyme
1277 Textus chondroideus Chondroid tissue Chondroid tissue
1278 Centrum ossificationis Ossification centre Ossification centre
1279 Motus detondens Shearing movement Shearing movement
1280 Stratum osteoblasticum Osteoblastic layer Osteoblastic layer
1281 Osteoblastus Osteoblast Osteoblast
1 Ejaculatio The reflex process of ejaculation occurs in two phases: in the first – emission – contraction of smooth muscle of glands and ducts delivers the various components of semen into the prostatic urethra;
in the second – ejaculation proper – the striated muscles of the urogenital triangle (particularly the bulbospongiosus muscles) contract spasmodically and expel semen from the urethra.
2 Embryogenesis Embryogenesis is the process of embryo formation. It entails the formation of the principal organs and systems and the acquisition of uniquely human surface features that are apparent with the
unaided eye. The process begins at fertilization and ends, somewhat arbitrarily, 56 days later.
3 Fetogenesis Fetogenesis entails the growth and differentiation, particularly functional differentiation, of the conceptus after embryogenesis is completed. It thus begins on day 57, following the Stage 23 embryo
that already has its principal organs, systems and distinctly human features, and ends at birth, when the fetus becomes a newborn infant or neonate. The time in which fetogenesis occurs may be divided into early,
intermediate and late fetal periods, which correspond to the trimesters of pregnancy in which they occur. There is, however, no agreement on precisely which weeks are encompassed by the first trimester; here it is
regarded as beginning at fertilization and as consisting of embryogenesis and the early fetal period, the 9th to the 13th post-fertilization weeks.
4 Cyclus vaginalis Cyclical changes in the stratified squamous epithelium of the vagina are not obvious in histological sections: under normal circumstances its desquamated cells remain nucleated and it does not
keratinize. However, according to Papanicolaou, there is a relative increase in acidophilic cells with small dark nuclei at the time of ovulation and thus three phases of the vaginal cycle may be recognized
(Papanicolaou GN. The sexual cycle in the human female as revealed by vaginal smears. Am J Anat 1933;52:519-637). The changes at ovulation may represent a prekeratinization process, which is completed
when the epithelium is exposed to the air, as in cases of prolapse.
5 Conceptus Conceptus refers to the entire product of conception from fertilization onwards.
6 Cyema The cyema is the embryonic or fetal part of the conceptus and thus excludes the developmental adnexa (q.v.), which are the placenta, umbilical cord and extra-embryonic membranes (O’Rahilly R, Müller,
F. Human Embryology and Teratology, 3rd ed. New York: Wiley-Liss; 2001).
7 Embryo [St.1 ad 23] Both embryonic and extra-embryonic cell lineages extend forward from the zygote and both extra-embryonic and embryonic tissues are necessary for normal development. Nevertheless, it
has been argued that to include the early stages in the use of the term embryo is misleading because a discrete and identifiable population of exclusively embryonic or cyemic cells does not exist until gastrulation is
under way and because most of the tissues formed prior to this are extra-embryonic or adnexal (Johnson MH and Selwood L. Nomenclature of early development in mammals. Reprod Fertil Dev 1996;8:759-64).
However, communication risks attend the redefining of a commonly and colloquially used term such as embryo and scientific purposes can be as well served by not redefining the term but defining the cells present
at a particular time. Embryo remains the preferred term for all 23 Carnegie Stages.
8 Adnexa developmentalia; Membranae embryonicae et fetales; Adnexa embryonica et fetalia The Latin word adnexum has been used almost exclusively in this plural form to indicate the structures (more than
one) adjacent to or subservient to a major structure. The form adnexae, although frequently used, is incorrect. The developmental adnexa are commonly referred to as the 'fetal membranes'. This is inaccurate
since they include the trophoblast, amnion, chroin, umbilical vesicle, allantoic vesicle, placenta and umbilical cord (O'Rahilly R, Muller F. Human Embryology and Teratology. 3rd ed. New York: Wiley-Liss; 2001).
9 Phasis pregastrulationis The pregastrulation and postgastrulation phases of the embryonic period and the fetal period are stages of prenatal development, each with its own distinctive characteristics, particularly
in respect of its responses to teratogens. The pregastrulation phase begins at fertilization, continues through cleavage and implantation and ends with the establishment of a definite primitive streak in Carnegie
Stage 6b at about two and a half weeks. It is a phase characterized by rapid increase in cell numbers and by regulation. As a result, response to teratogens is uncertain: induced errors of development may regulate
but, if they do not, the errors are likely to be of such magnitude that early spontaneous abortion follows.
10 Phasis preparatoria; Phasis embryogenica The characteristic of the preparative phase is that it is spent preparing extra-embryonic membranes and presumptive embryonic cells but that no cells of the
conceptus have yet been determined as substantive embryonic cells. It has therefore been called the embryogenic phase (Johnson MH and Selwood L. Nomenclature of early development in mammals. Reprod
Fertil Dev. 1996;8:759-64). The term “pre-embryonic stage”, which has been used in legal and clinical contexts, is not recommended.
11 Pregnatio cervicalis; Pregnatio ectopica; Pregnatio extrauterina Although appropriately listed with uterine pregnancy sites, a cervical pregnancy is often considered to be an ectopic pregnancy.
13 Aetas a fecundiatione Fertilization age begins at the time of fertilization with the sperm penetrating the oocyte and the formation of the zygote. It is the true age of the conceptus and the preferred measure.
14 Aetas ab ovulatione Ovulation age begins on the day of the ovulation that preceded fertilization and the formation of the zygote: it is about 0.5 day longer than fertilization age.
15 Aetas ab inseminatione Insemination age begins when the sperm and oocyte are introduced in artificial insemination or in vitro fertilization.
16 Hebdomades post coitum Coital weeks begin from the time of the coitus that resulted in the pregnancy. Normally, fertilization occurs early in the first coital week. Since the embryo does not exist for the first part
of the first coital week, the term coital age is inappropriate.
17 Hebdomades post menses ultimas Menstrual ("gestational") weeks begin from the first day of the mother's last menstrual period [LMP] before becoming pregnant and are the usual measure in obstetric practice.
Since the embryo does not usually come into being until the first two menstrual weeks have passed, the term menstrual “age” is inappropriate. The term gestational age is superfluous, ambiguous and should be
abandoned, it having been variously equated with menstrual weeks, ovulation age and fertilization age (O’Rahilly R, Müller F. Developmental Stages in Human Embryos: Revised and New Measurements. Cells
Tissues Organs 2010;192:73-84).
18 Longitudo maxima Greatest length [GL] is the preferred measure of length, being independent of fixed points, which are not always easy to determine. GL coincides with crown-rump length [CRL] at Stages 11
and 12; GL is generally more than CRL and coincides with neck-rump length from Stages 13-17; GL and CRL again coincide from Stages 18-20 onwards (O’Rahilly R, Müller F. Developmental Stages in Human
Embryos: Revised and New Measurements. Cells Tissues Organs 2010;192:73-84).
19 Gemini conjuncti See Spencer R. Conjoined twins. Baltimore: Johns Hopkins University Press; 2003. In conjoined twins, as elsewhere, convention has the suffixes –ia in Latin and –y in English indicating the
condition; the suffix –us, in either language, refers to an individual with that condition.
20 Genum a parente impressum Genomic imprinting occurs during meiosis II of gametogenesis and persists until the primary gametocyte stage in the next generation.
21 Polus animalis Being microlecithal, the human primary oocyte does not exhibit the obvious polarity characteristic of more richly yolked oocytes. It does, however, exhibit some degree of asymmetry in
distribution of cytoplasmic elements. The animal pole of the arrested secondary oocyte is identified by the position of the second meoitic spindle and the lack of microvilli on the cell membrane overlying it. After
fertilization, the animal pole of the ootid is characterized by the presence of the female and male pronuclei. There is no necessary relationship between the animal-vegetal axis and the future embryonic-
abembryonic (dorsoventral) axis. In some (but not all) mouse zygotes, the animal-vegetal axis corresponds to the long axis of the ellipsoid blastocyst and thus to the anteroposterior axis of the embryo. In these
cases the animal-vegetal axis is orthogonal to the embryonic-abembryonic axis (Selwood L, Johnson MH. Trophoblast and hypoblast in the monotreme, marsupial and eutherian mammal: evolution and origins.
BioEssays 2006;28:128-145).
22 Pellucidagenesis; Zonagenesis Although the term zonagenesis is widely used in zoology, it is not recommended as it lacks a locational adjective and could thus apply to any zone. Although the zona pellucida
cannot be seen with the light microscope before the primary ovarian follicle has developed, the heavily glycosylated proteins ZP 1-3 can be demonstrated in the oocytes and follicle cells of primordial follicles (Gook
DA, Edgar DH, Borg J and Martic M. Detection of zona pellucida proteins during human folliculogenesis. Hum Reprod 2008;23:394-402).
23 Regulatio The process by which the developmental fates or rates of development of cells of embryonic subsystems may change during embryonic development, thereby permitting normal integrated
development of the embryo as a whole and compensating for anomalies. It is the result of changes in gene expression; moreover, since the DNA sequence that comprises the genome remains unchanged during
the differentiation of systems, organs, tissues and cell-types, regulation is said to be an epigenetic process. The human zygote is said to be regulatory because in it the primordia of tissues and organs are not
24 Differentiatio cellularum in zygoto findenti Differentiation of the cells of the cleaving zygote into outer blastomeres, which are polarized, and inner blastomeres, which are not.
25 Polarisatio cellularum externarum Transformation of rounded, radially symmetrical outer blastomeres into highly asymmetric cells with the characteristics of epithelia. Longitudinal divisions of polarized cells are
conservative, resulting only in more polarized cells. Transverse divisions of polarized cells are differentiative, resulting in both embryoblastic cells and polarized cells. Cells remaining polarized give rise to
trophoblast (Johnson MH. Origin of pluriblast and trophoblast in the eutherian conceptus. Reprod Fertil Dev 1996;8:699-709).
26 Divisio differentiativa embryoblasti; Divisio differentiativa massae cellularis internae; Divisio differentiativa pluriblasti Differentiation of the cells of the embryoblast, inner cell mass or pluriblast into the dorsal cells
of the epiblast and the ventral cells of the hypoblast, with a basal lamina between them.
27 Morphogenesis The development of shape, size or other feature of a particular organ or of a part or the whole of the body. “The word ‘morphogenesis’ is often used in a broad sense to refer to many aspects of
development, but when used strictly it should mean the moulding of cells and tissues into definite shapes” (Waddington CH. Principles of Embryology. London: George Allan & Unwin; 1956). In this strict sense, it
refers particularly to the wide-ranging phenomena associated with gastrulation and organogenesis and to local phenomena like budding, branching and clefting (Hogan BLM. Morphogenesis. Cell 1999;96:225-233).
28 Blastema An identifiable mass of rapidly proliferating undifferentiated cells that gives rise to a differentiated structure/organ.
29 Primordium A term applied to a structure making its first appearance as a differentiating structure. Anlage, from the German, is a synonym. It is now appreciated that, particularly in branching morphogenesis,
an epithelial primordium may be preceded by a mesenchymal primordium, which determines the pattern of arborization (Denny PC, Ball WD, Redman RS. Salivary glands: a paradigm for diversity of gland
development. Crit Rev Biol Med 1997;8:51-75).
30 Rudimentum; Vestigium These terms are not interchangeable: a rudiment (from the Latin rudimentum – that which is unwrought) is an underdeveloped or immature part or organ; a vestige (from the Latin
vestigium – that which is tracked) is a part or organ that has become reduced in function and/or size in the course of phylogeny; some vestiges, nevertheless, play an important part in ontogeny.
31 Status presumptivus The condition of a tissue, region or organ that will, in the course of normal development, become a morphologically differentiated tissue, region or organ. A structure may be presumptive
solely by virtue of its position or it may have undergone determination or chemodifferentiation but as yet shows no visible signs of differentiation.
33 Formatio ansae See, for example: Männer J. The anatomy of cardiac looping: a step towards the understanding of the morphogenesis of several forms of congenital heart malformations (Clin Anat 2009;22:21-
35).
34 Formatio primaria corporis Primary body development involves the primary germ layers and neural plate more or less directly. It includes primary neurulation, the formation of somites 1-29, of spinal ganglia 1-
25, of the foregut, midgut and hindgut and of the corresponding part of the notochord.
35 Formatio secundaria corporis Secondary body development does not involve the germ layers: in it structures develop directly from the axial dense mesenchyme of the caudal eminence or tail bud, which is the
remnant of the primitive streak. It includes secondary neurulation, the formation of somites 30-39, of spinal ganglia 26-35, of the most caudal gut and of the corresponding part of the notochord.
36 Gastrulatio It has been said that the term gastrulation is inappropriate as it refers to the invagination of a monolayered blastula to form a bilayered gastrula containing an endoderm-lined archenteron (O'Rahilly
R and Müller F. Human embryology and teratology. 3rd ed. New York: Wylie-Liss; 2001). While this was the original meaning of gastrulation, its meaning has undergone a profound change (Collins P and Billett FS.
The terminology of early development: history, concepts, and current usage. Clin Anat 1995;8:418-25). It may now be defined as the formative process by which the three germ layers and an axial organization are
37 Inflatio The ballooning model has succeeded the segmental model of heart chamber formation (Horsthuis T, Christoffels VM, Anderson RH, Moorman AFM. Can recent insights into cardiac development
improve our understanding of congenitally malformed hearts? Clin Anat 2009;22:4-20).
38 Invectio Introduction of material into a structure from outside. An example is the process by which material from the mesocardium is added to the venous and arterial poles of the early heart tube (Horsthuis T,
Christoffels VM, Anderson RH, Moorman AFM. Can recent insights into cardiac development improve our understanding of congenitally malformed hearts? Clin Anat 2009;22:4-20).
39 Maturatio Maturation may be defined as the progressive acquisition of definitive structure and function: its prenatal aspects, particularly, are within the compass of Terminologia Embryologica.
40 Morphogenesis gemmans Budding morphogenesis and the ensuing canalization have been most widely studied in the lung, kidney, mammary gland and salivary glands and the processes detailed for it are
generally followed elsewhere (Varner VD, Nelson CM. Cellular and physical mechanisms of branching morphogenesis. Development 2014;141:2750-2759).
41 Morphogenesis ramificans Branching morphogenesis is the process of forming organized patterns of epithelial cords and then tubules in organs such as the kidney, glands and lungs. It appears to be
determined by mesenchyme and regulated by a wide range of factors (Williams MJ, Clark P. Microscopic analysis of the cellular events during scatter factor/hepatocyte growth factor-induced epithelial
tubulogenesis. J Anat 2003;203:483-503). The term tubulogenesis is not recommended in this context as the product is initially solid and only canalizes later.
42 Morphogenesis findens Clefting is the process in which a terminal bud is cleaved into multiple lobules with the ingrowth of mesenchyme and the deposition of extracellular matrix. In some organs, such as the
lung, budding, branching and clefting each occur at different stages of development, whereas in salivary glands clefting appears to predominate (Hogan BLM. Morphogenesis. Cell 1999;96:225-233).
43 Motus condensationis; Motus densationis Movement occurring in a morphogenetic field, called a densation field (Blechschmidt E, Gasser R. Biokinetics and biodynamics of human differentiation. Springfield:
Charles C Thomas; 1978). With loss of intercellular fluid, the cells come closer together. The first appearance of the skeleton is as mesenchymal condensations. The cells have rather spherical cell bodies and very
little intercellular substance present between them. They show no particular orientation which means that they are under tension stresses that are equal in all directions. A densation field is characterized by its
position.
44 Motus dilatationis Movement occurring in a morphogenetic field, called a dilation field (Blechschmidt E, Gasser R. Biokinetics and biodynamics of human differentiation. Springfield: Charles C Thomas; 1978).
The field is spatially and kinetically highly organized. Dilation occurs in mesenchymal tissue when it becomes elongated and slenderized by pull in a particular direction without transverse compression. The
primordia of skeletal, cardiac and smooth muscle fibres and fibre systems arise in dilation fields. The shape of a muscle is closely related to its position while its structure is closely related to its shape.
45 Motus expansionis longitudinalis; Motus distusionalis Movement occurring in a morphogenetic field, called a distusion field (Blechschmidt E, Gasser R. Biokinetics and biodynamics of human differentiation.
Springfield: Charles C Thomas; 1978). The spherical mesenchymal cells of a densation field become compressed by opposing forces in the longitudinal axis of the field. Initially, such zones of flattened cells are
located only in the centre of a field that has attained sufficient size. For example, cells in such a contusion field become disc-shaped and develop into cartilage cells.
46 Motio involutionis; Involutio The rolling-in of cells over a rim. In this context the term involutionary movement is preferred because of the different connotations of the term involution.
47 Motus translationis; Migratio When re-examined appropriately (Gasser RF. Evidence that some events in mammalian embryogenesis can result from differential growth, making migration unnecessary. Anat
Rec B New Anat.2006;289B:53-63), many examples of changing positional relations turn out not to be examples of true migration.Cell migration may be false or true. In false migration, structures do not move from
one site to another but their positional relationships change and they become separated as an embryo enlarges and changes shape. See, for example: Freeman B. The active migration of germ cells in the embryos
of mice and men is a myth (Reproduction 2003;125:635-643), which explains the relocation of primordial germ cells from the wall of the umbilical vesicle to that of the hindgut by growth movements and shape
changes. True migration, such as occurs in cytokinesisis in the cerebellum and the cerebral cortex, is the movement of cells, cell groups and organs from one site to another, among, around, through or over other
structures, in relation to a central reference point that moves minimally as the embryo enlarges or changes shape.
49 Neurulatio secundaria Secondary neurulation is the process entailing canalization that leads to the formation of the spinal cord beyond the limits of primary neurulation. It occurs after the closure of the caudal
neuropore in Stage 12, and thus in Stages 13-18. Axial dense mesenchyme in the caudal eminence forms a neural cord in continuity with the neural tube: the cavity in the neural tube extends into the neural cord
(Müller F, O'Rahilly R. The development of the human brain, the closure of the caudal neuropore, and the beginning of secondary neurulation at stage 12. Anat Embryol 1987;176:413-30). The term secondary
neurulation is best confined to the process rather than applied to the phase of development in which it occurs, which is best referred to as secondary body development.
50 Transitio epitheliomesenchymalis For sessile cells to become free and migrate they must undergo an epitheliomesenchymal transition, first becoming flask-shaped, with the dissolution of juxtaluminal junctions,
and then becoming frankly mesenchymal.
51 Transitio mesenchymoepithelialis Mesenchyme cells that have reached their destinations may condense and revert to sessile epithelial cells, by polarizing, developing basal laminae and specialized
juxtaluminal junctions. Some such epithelia may subsequently undergo an epitheliomesenchymal transition.
52 Ontogenesis Ontogenesis is defined here as the development of the individual, beginning at fertilization and ending at death. It thus covers the principal concerns of this terminology (embryogenesis,
fetogenesis and immediate postnatal development) but extends beyond them.
53 Tempus fetale The pregastrulation and postgastrulation phases of the embryonic period and the fetal period are stages of prenatal development, each with their own distinctive characteristics, particularly in
respect of their responses to teratogens. The fetal period is taken, somewhat arbitrarily, to begin on day 57, by which time the embryo has already acquired the distinctly human features that are apparent with the
unaided eye, and ends at birth, when the fetus becomes a newborn infant or neonate. The main events of organ formation having been completed by Carnegie Stage 23, the fetal period is mainly one of growth and
differentiation, particularly functional differentiation, in preparation for extra-uterine life. Notable in this context is the skeletal system, in which cartilaginous precursors are being replaced by bone, and the nervous
system, which is forming functional connections. Thus anomalies arising during this period entail disturbances of growth, of hard tissues and of neural connections.
54 Aetas fetalis Fetal age is usually given in weeks and is determined by the use of various starting points, of which only fertilization, insemination and ovulation are valid.
55 Tempus fetale initiale; fetus hebdomadis nonae ad hebdomadam tertiam decimam The early fetal period here corresponds to that of the 9 to 13 week fetus and ends at 90 days and about 90mm: the conclusion
of the “second sous-stade de finition histogénétique, de réglage des proportions” (Guyot R. Théorie nouvelle sur les âges de la vie. 2nd ed. Paris: Barré & Dayez; 1985) and probably the end of the first trimester.
There is, however, no agreement on precisely which weeks are encompassed by the first trimester. The attributes of fetuses have not been subjected to the same systematic, intensive investigation as have
embryos. Nevertheless, there are attributes other than size and weight that characterize progression during the early fetal period.
56 Tempus fetale intermedium; Fetus trimestris secundi The intermediate fetal period here corresponds to that of the second trimester fetus and thus begins at 90 days and about 90mm, after the conclusion of the
“second sous-stade de finition histogénétique, de réglage des proportions” (Guyot R. Théorie nouvelle sur les âges de la vie. 2nd ed. Paris: Barré & Dayez; 1985).
57 Tempus fetale serum; Fetus trimestri tertii The late fetal period corresponds to that of the third trimester fetus.
58 Tempus perinatale The perinatal period extends from immediately prior to birth, through birth and through the first 7 days of postnatal life, the early neonatal period.
59 Tempus neonatale The first 7 days after birth constitute the early neonatal period. The following 21 days of postnatal life constitute the late neonatal period, which thus ends with day 28.
61 Paedomorphosis Paedomorphosis is exhibited in features such as the human flat face, position of the foramen magnum, retarded skeletal development and continuation of fetal growth rates into infancy and
childhood.
62 Fertilisatio post penetrationem spermatozoi Some of the features included in this section are inferred because they are common to all mammalian fertilization: others, such as Zonal reaction and Fertilization
cone, have been observed in the human as a result of in vitro fertilization.
63 Ootidium; Ovum; Embryo pronuclearis [Gradus Ib] The imprecise term ovum has been variously applied, alone or qualified, to stages from the primary oocyte to the implanting blastocyst and beyond. The use of
the more precise term is recommended. In mammals, it is the secondary oocyte, arrested in the metaphase of meiosis II, which is penetrated and thus best referred to as a penetrated oocyte until meiosis II has
been completed. Penetration activates the oocyte into completing meiosis II, with the formation of the ootid and the second polar body. The mammalian ootid contains two separate haploid elements, the female and
male pronuclei. As these two elements fuse into a single diploid aggregation of chromosomes, the ootid becomes a zygote.
64 Corpus polare secundum; Polocytus secundarius A line through the centres of the zygote and the second polar body defines the polar axis and indicates the plane of the first cleavage division (Veeck L L,
Zaninovic N. An atlas of human blastocysts. New York: Parthenon Publishing Group; 2003).
65 Numerus diploideus chromosomatum nonreplicatorum; Numerus diploideus chromosomatum nonreplicatorum This relates to the condition in the penetrated (or definitive) oocyte [2N].
66 Syngamia Traditionally, syngamy has meant sexual reproduction or, more specifically, the fusion of gametes. However, in in vitro fertilization it has come to describe a stage, beginning some 21-32 hr after
insemination, in which maternal and paternal chromosomes intermingle, although this is not easily discernible by ordinary microscopy (Sathananthan H, Trounson AO, Wood C. Atlas of fine structure of human
sperm penetration, eggs and embryos cultured in vitro. New York: Praeger Publishers; 1986).
67 Activatio prima genorum zygoticorum This first transcription produces only a minor population of mRNAs whereas the second transcription (ZGA2), in the two-celled embryo, produces a major population: most
maternal mRNA is degraded at this time although maternal proteins persist into the blastocyst stage (Selwood L, Johnson MH. Trophoblast and hypoblast in the monotreme, marsupial and eutherian mammal:
evolution and origins. BioEssays 2006;28:128-145); De Paepe C, Krivega M, Cauffman G, Geens M, van de Velde H. Totipotency and lineage segregation in the human embryo. Mol Hum Reprod 2014;20:599-618).
68 Tempus embryonicum; Gradus carnegiensis [1-23] There are 23 defined stages of development during the embryonic period, which begins at fertilization and ends, somewhat arbitrarily, 56 days later, by which
time the embryo has already acquired uniquely human surface features that are apparent with the unaided eye. The stages are the internationally accepted Carnegie Stages (O'Rahilly R, Müller F. Developmental
stages in human embryos. Washington DC: Carnegie Institution of Washington; 1987). Each Carnegie Stage is an arbitrarily defined cut through the time axis of the embryo and is based upon carefully-defined
external and internal morphological criteria and not length or age. Thus, embryos of a particular length or age are not necessarily embryos of a particular stage. Carnegie Stage cannot be assigned solely on the
basis of such measurements. See Dickey RP, Gasser RF. Ulltrasound evidence for variability in the size and development of normal human embryos before the tenth postinsemination week after assisted
reproductive technologies. Hum Reprod 1993;8:331-337; Wisser J, Dirschedl P, Krone S. Estimation of gestational age by transvaginal sonographic measurement of greatest embryonic length in dated human
embryos. Ultrasound Obstet Gynecol 1994;4:457-462; Blaas HG-K, Eik-Nes SH, Kiserud T, Hellevik LR. Early development of the forebrain and midbrain: A longitudinal ultrasound study from 7 to 12 weeks of
gestation. Ultrasound Obstetr Gynecol 1994;16:25-29; Pooh RK, Kurjak A, eds Fetal Neurology. Jaypee, St.Louis; 2009; Pooh RK, Shiota K, Kurjak A. Imaging of the human embryo with magnetic resonance
imaging microscopy and high-resolution transvaginal 3-dimensional sonography: Human embryology in the 21st century. Am J Obstet Gynecol 2011;204:77.e1-77.e16.). See also O’Rahilly and Müller F, op.cit. for
further discussion of staging and for further references on ultrasound studies. Data have also been derived from MR studies on embryos from the Kyoto Collection of Embryos: Yamada S, Samtani RR, Lee ES,
Lockett E, Uwabe C, Shiota K, et al. Developmental atlas of the early first trimester human embryo. Dev Dyn 2010; 239:1585-1595.
69 Embryo pregastrulationis [St.1 ad 6a] The term pregastrulation embryo is useful because such an embryo has distinctive attributes. The foreshortened term “pre-embryo”, which has been used in legal and
clinical contexts, is not recommended.
70 Spatium subzonale; Spatium subcapsulare The commonly used term perivitelline space is inappropriate for the space surrounding the human oocyte, which is deficient in yolk (Latin - vitellus).
72 Degradatio mitochondriorum paternalium See Sutovsky P, Van Leyen K, McCauley T, Day BN, Sutovsky M. Degradation of paternal mitochondria after fertilization: implications for heteroplasmy, assisted
reproductive technologies and mtDNA inheritance. Department of Animal Science, University of Missouri-Columbia, MO, USA. [email protected], Reprod Biomed Online. 2004 Jan;8(1):24-33.
73 Intercorpus The interbody is a prominent intracytoplasmic electron-dense contractile structure in the equatorial plane of the second meiotic spindle, extending from the penetrated oocyte into the extruding
second polar body. Fine electron-dense particles of unknown chemical nature are associated with spindle microtubules and the interbody represents the site of detachment of the second polar body and
reconstitution of the cell membranes of the embryo and second polar body.
74 Zygotum findens [Gradus II] Embryos of Carnegie Stage 2 consist of between 2 and about 32 cells but have no blastocystic cavity by light microscopy. They are generally 0.1-0.2mm in diameter and about 2-3
days old.
75 Morula Stage 2 embryos from 12 to about 32 cells and without a blastocystic cavity are called morulae (from Latin Morus, mulberry). The term is not ideal because, unlike the amphibian morula, for which the
term was coined, the human morula gives rise to extra-embryonic as well as embryonic tissues. Nevertheless, when the number of blastomeres cannot be counted, there is no other term to describe the solid mass
that precedes the formation of the blastocystic cavity.
76 Cellula externa morulae; Cellula trophoblastica presumptiva; Cellula polarisata; Polarblastus The outer cells of the morula are polarized and are asymmetrical cells with the characteristics of epithelia. Their
longitudinal divisions are conservative and result only in more polarized cells. Their transverse divisions are differentiative and result in both unpolarized embryoblastic cells and polarized cells. Cells remaining
polarized become trophoblast (Johnson MH. Origin of pluriblast and trophoblast in the eutherian conceptus. Reprod Fertil Dev 1996;8:699-709). The term polarblast appropriately describes the tissue.
77 Cellula interna morulae; Cellula embryoblastica presumptiva; Pluriblastus initialis The inner cells of the morula are unpolarized and remain rounded and radially symmetrical. Their divisions are conservative and
result only in more unpolarized cells. They will become the embryoblast or inner cell mass. The term pluriblast (Johnson MH. Origin of pluriblast and trophoblast in the eutherian conceptus. Reprod Fertil Dev
1996;8:699-709) recognizes the fact that its derivatives are both extra-embryonic or adnexal and embryonic or cyemic.
78 Blastocystis libera [Gradus III] Embryos of Carnegie Stage 3 are free blastocysts with a blastocystic cavity by light microscopy. They consist of up to 90 cells, of which about 30 are inner cell mass cells. They
are about 4-5 days old and are generally 0.1-0.2mm in diameter.
79 Embryoblastus; Massa cellularis interna; Pluriblastus serior The term embryoblast is widely used although the derivatives of this tissue are both extra-embryonic or adnexal and embryonic or cyemic. The term
inner cell mass is also used but cannot be used as a comparative term as the corresponding cells are not inner in many, if not all, marsupials (Johnson MH and Selwood L. Nomenclature of early development in
mammals. Reprod Fertil Dev 1996;8:759-64). The term pluriblast has neither of these disadvantages.
80 Epithelium primordiale The tissues of the pre-implantation embryo proper are all epithelial in that their cells are sessile, are polarized between a free surface and a basal lamina and they have specialized
juxtaluminal junctions and little intercellular substance. Daughter cells may be epithelial or, as a result of epitheliomesenchymal transition, become mesenchymal (see below).
81 Epiblastus The term epiblast is preferred for this tissue as it provides appropriate information about its site, fate and potential. Primary ectoderm is less preferred both because the term has been used to include
amnioblast and because current usage postpones the use of the suffix -derm until after gastrulation (Johnson MH and Selwood L. Nomenclature of early development in mammals. Reprod Fertil Dev 1996;8:759-
64). Alternatives that include the term ectoderm, the use of which should be limited to the cells remaining on the dorsal surface of the embryo after the early somite stage, are not recommended.
82 Hypoblastus The term hypoblast is preferred for this tissue as it provides appropriate information about its site, fate and potential. It appears to induce the formation of the primordial amniotic cavity
(Coucouvanis E, Martin GR. Signals for death and survival: a two-step mechanism for cavitation in the vertebrate embryo. Cell 1995;83:279-287) and of axial patterning in the epiblast, including the forebrain
(Beddington RSP, Robertson EP. Axis development and early asymmetry in mammals. Cell 1999;96:195-209). The term primary endoderm is less preferred both because the term has been used to include the
83 Mesenchyma A tissue consisting of free cells without polarity or specialized juxtaluminal junctions. The loosely arranged, often stellate, cells are suspended in a gelatinous matrix and are amoeboid and actively
phagocytic. As they migrate their processes make temporary contact with each other, with overlying epithelial cells and with their basal laminae. The outcome of eptheliomesenchymal transition is that there are two
types of tissue, namely, epithelial and mesenchymal. These tissue types do not correspond to the primary germ layers, ectoderm, endoderm and mesoderm [q.v.].
84 Cavitas blastocystica The term blastocystic cavity is recommended, rather than blastocoel, because the cavity is not homologous with the blastocoel of amphibians and birds, the homologue of which is "the
very narrow slit confined between the epiblast and hypoblast" (Eyal-Giladi H. Establishment of the axis in chordates: facts and speculations. Development 1997;124:2285-2296).
85 Trophoblastus; Trophoectoderma The term trophoblast is preferred for this tissue, which is defined as the earliest appearing stem cell population dedicated to nourishment of future embryonic tissues. Its cells
are adhesive, migratory and, initially, multinucleate. They appear, at least in the mouse, to signal the specification of primordial germ cells and the allantois in the caudal epiblast (Lawson KA, Dunn NR, Roelen BA,
Zeinstra LM, Davies AM, Wright CV, Corving JP, Hogan BL. Bmp4 is required for the generation of primordial germ cells in the mouse embryo. Genes Dev 1999;13:424-436). The term trophectoderm and its
variants are not recommended because current usage postpones the use of the suffix -derm until after gastrulation (Johnson MH and Selwood L. Nomenclature of early development in mammals. Reprod Fertil Dev
1996;8:759-64). Alternatives that include the term ectoderm, the use of which should be limited to the cells remaining on the dorsal surface of the embryo after the early somite stage, are not recommended.
86 Blastocystis unilaminaris [Gradus III] The cavity of a unilaminar blastocyst is surrounded by a single layered extra-embryonic ectodermal membrane, the trophoblast; in the bilaminar blastocyst, the trophoblast
is lined by extra-embryonic endoblast; the interposition of extra-embryonic mesoderm creates the trilaminar blastocyst.
87 Insignia miscellanea cellularum trophoblastorum nondifferentiatarum These miscellaneous features have only been observed in vitro. Similar observations on the differentiated trophoblast are not readily
available.
88 Blastocystis adhaerens [Gradus IV] An embryo of Carnegie Stage 4 is an attaching blastocyst but no such in vivo human specimen has been recorded. Previously reliance was placed on those of the macaque
(Heuser CH, Streeter GL. Development of the macaque embryo. Contrib Embryol 1941;29:15-55) but direct information about Stage 4 is now derived from in vitro experiments in which blastocysts are placed on
monolayers of cultured endometrial epithelial cells. Embryos of Stage 4 would be about 6 days old and about 0.1-0.2mm in diameter.
89 Cavitas amniotica primordialis It appears that a primordial amniotic cavity forms by cavitation within the epiblast, that the roof of the primordial amniotic cavity breaks down creating a transient tropho-epiblastic
cavity and that the definitive amniotic cavity becomes roofed by cells that arise from the margins of the epiblast. There is no primordial amniotic cavity in embryos of Stage 3 and while most embryos of Stage 5a
have a tropho-epiblastic cavity, the “earliest known human implantation stage” (Carnegie No 8020) has a primordial amniotic cavity (Luckett WP.The development of primordial and definitive amniotic cavities in early
Rhesus monkey and human embryos. Am J Anat 1975;144:149-168). It is therefore presumed that for most embryos a primordial amniotic cavity occurs in Stage 4.
90 Blastocystis implantata; Blastocystis invadens; Conceptus previllosus [St.5] Embryos of Carnegie Stage 5 are implanted but previllous blastocysts. Their trophoblast is solid in Stage 5a, contains isolated
lacunae in Stage 5b and contains intercommunicating lacunae in Stage 5c. The embryonic disc in embryos of Stage 5, which are about 7-12 days old, is generally 0.1-0.2mm in diameter.
91 Margo syncytiodecidualis The endometrium responds to the presence of syncytiotrophoblast by undergoing the predecidual reaction, characterized by oedema and then saw–toothed glands, particularly in the
stratum spongiosum. It is thenceforward called decidua.
92 Cavitas trophoepiblastica The trophoepiblastic cavity is apparently formed by the breakdown of the roof of the primordial amniotic cavity. See above.
93 Amnioblastus; Cellulae amniogenicae; Ectoderma amnioticum The term amnioblast is preferred for this tissue as it provides appropriate information about its site, fate and potential. Extra-embryonic ectoderm is
least preferred because current usage postpones the use of the suffix -derm until after gastrulation (Johnson MH and Selwood L. Nomenclature of early development in mammals. Reprod Fertil Dev 1996;8:759-64).
Alternatives that include the suffix -derm are not recommended.
95 Area caudalis mesoblastogenica An area at the future caudal margin of the epiblast, which precedes the appearance of a definite primitive streak [St.6b]. Extra-embryonic mesoblast is thought to arise from this
area rather than by delamination from the trophoblast.
96 Mesoblastus extraembryonicus The term mesoblast is preferred for this tissue because it provides appropriate information on its site, fate and potential. Lankester used the term to describe those cells, which
he thought derived from enteric cells (extra-embryonic endoblast), separated, spread out, became amoebiform and “crawled all over the inner wall of the ectodermic vesicle (blastocoele or blastocystic cavity)
(Lankester R. Notes on the embryology and classification of the animal kingdom. Q J Microscop Sci 1877;17:399-454). A term is necessary to describe the tissues found outside the extra-embryonic endoblast and
inside the trophoblast before gastrulation and the use of the suffix -blast is a corollary of current usage (Johnson MH, Selwood L. Nomenclature of early development in mammals. Reprod Fertil Dev 1996;8:759-64).
The qualifying adjective extra-embryonic is necessary because the term mesoblast has been used to describe the free cells that migrate between the epiblast and intra-embryonic endoderm (Collins P, Billett FS.
The terminology of early development: history, concepts, and current usage. Clin Anat 1995;8:418-25) and to denote a temporary, embryonic cell lineage, which will later generate either an epithelial or a free-cell
arrangement (Collins P. Embryology and development. In: Williams PL, Bannister LH, Berry MM, Collins P, Dyson M, Dussek J, Ferguson MWJ, editors. Gray’s Anatomy 38th ed. Edinburgh: Churchill Livingstone;
1995).
97 Textus angioblasticus mesoblasti / Crista previllosa mesoblasti Extra-embryonic mesoblastic tissues produced from the epiblast before gastrulation.
98 Reticulum extraembryonicum; Magma reticulare Extra-embryonic mesoblast produced, initially from the hypoblast, before gastrulation. At least in the rhesus monkey, the cells of both the hypoblast and the
initial reticulum are mitotically active (Enders AC, King BF. Formation and differentiation of extraembryonic mesoderm in the rhesus monkey. Am J Anat 1988;181:327-340) so that later reticulum may have arisen
from either source.
99 Endoblastus extraembryonicus; Membrana exocoelomica The term endoblast is preferred for this tissue as it provides appropriate information about its site, fate and potential. The term primary endoderm is
least preferred both because it applies also to the hypoblast and also because current usage postpones the use of the suffix -derm until after gastrulation (Johnson MH, Selwood L. Nomenclature of early
development in mammals. Reprod Fertil Dev 1996;8:759-64). Other alternatives that include the term endoderm are not recommended.
100 Vesicula umbilicalis primaria / Saccus vitellinus primarius; Cavitas vesiculae umbilicalis primariae / Cavitas sacci vitellini primarii; Vesicula umbilicalis secundaria / Saccus vitellinus secundarius; Cavitas
vesiculae umbilicalis secundariae / Cavitas sacci vitellini secundarii The term umbilical vesicle, which has been in use for many years, is preferred because yolk (Latin vitellus) is not present in the human vesicle
and because the term indicates location, the vesicle being a feature of the umbilical region of the embryo and becoming, at least partially, incorporated into the umbilical cord.
101 Circulus lacunosus vascularis The lacunar vascular circle is visible from the endometrial surface.
102 Lamina prechordalis precoqua “The first clear evidence of a (rostro)caudal embryonic axis appears [at Stage 5c] as a pronounced thickening of the [hypoblast] at the future cranial end of the embryonic disc to
form a prechordal plate.” (Luckett WP. Origin and differentiation of the yolk sac and extraembryonic mesoderm in presomite human and rhesus monkey embryos. Am J Anat 1979;152:59-98). However, this
thickening may not correspond to the prechordal plate but to the extra-embryonic rostral visceral endoderm or rostral marginal crescent found in other mammals (Viebahn C.The anterior margin of the mammalian
gastrula: comparative and phylogenetic aspects of its role in axis formation and head induction. Curr Top Dev Biol 1999;46:63-103). The prechordal plate proper may not appear until Stage 7.
103 Polus rostralis embryonis Although rostrum means a beak, a snout or the prow of a ship, rostral is commonly used as the opposite of caudal, particularly before the appearance of cephalic structures in Stage
8 or cranial structures in Stage 13, but also thereafter: it is also used in neuroanatomy to mean nearer the rostrum of the corpus callosum in the unfolded nervous system.
105 Latus dextrum embryonis Although the sides of the embryo can be recognized and cranial patterning occurs in Stage 5, it is not until Stage 6b, with the formation of the primitive node, that the molecular basis
for left-right asymmetry is established.
106 Conceptus villosus [Gradus VI] Embryos of Carnegie Stage 6 are villous conceptuses. There may be the suggestion of a primitive streak in embryos of Stage 6a or earlier, but one is definitely present in
embryos of Stage 6b. The embryonic discs of embryos of Stage 6 are generally about 0.2mm in diameter and they are about 17 days old.
107 Chorion frondosum The predecidual reaction around embryos of Carnegie Stage 5 becomes a full-blown decidual reaction around embryos of Carnegie Stage 6, with the transformation of stromal cells into
decidual cells: they become rounded or polyhedral and glycogen, lipids and mitochondria accumulate within their vacuolated cytoplasm.
108 Vasculogenesis Formation of a primordial capillary network from cells that differentiatie locally: they may be invading angioblasts (Risau W, Flamme I. Vasculogenesis. Annu Rev Cell Dev Biol. 1995:11;79-91)
or be endothelial progenitor cells (EPCs) which differentiate in splanchnopleuric mesenchyme or its derivatives.
109 Mesenchyma capitis Head mesenchyme is listed as present in embryos from Stage 6b onwards in the Edinburgh atlas and database of human developmental anatomy.
http://www.ana.ed.ac.uk/anatomy/database/humat/ but the primary source of the listing is not known. Its presence in embryos of Stage 7 may be inferred from Hill JP, Florian J. A young human embryo (embryo
Dobbin) with head-process and prechordal plate (Phil Trans Roy Soc London B 1931;219:443-486).
110 Linea primitive / Linea gastrulationis; Nodus primitivus / Nodus gastrulationis; Sulcus primitivus / Sulcus gastrulationis The terms primitive streak/node/groove are widely used but may be misuses of the term
primitive, which more usually refers to phylogeny rather than ontogeny. The alternative terms, gastrulation streak / node / groove, do not have this anomaly and, from a functional viewpoint, are more informative.
111 Mesoderma embryonicum; Mesoblastus These terms describe the intermediate germ layer of the trilaminar embryo, which will form bone, muscle and connective and blood-vascular tissues. Experimental
studies suggest that cells ingressing through the rimitive node and the rostral part of the primitive streak give rise to paraxial mesoderm and those through the middle part give rise to lateral plate mesoderm. The
qualifying adjective embryonic is necessary as long as terms such as primary mesoderm remain in use. The term mesoblast has been used (Collins P. Embryology and development. In: Williams PL, Bannister LH,
Berry MM, Collins P, Dyson M, Dussek J, Ferguson MWJ, editors. Gray’s Anatomy 38th ed. Edinburgh: Churchill Livingstone; 1995 and subsequent editions). However, the use of the suffix -derm for a germ layer
produced by gastrulation is preferred and has that restricted use here : the parts of the germ layer are paraxial and lateral plate mesoderms; their derivatives are either epithelial or mesenchymal and are named
accordingly.
112 Endoderma embryonicum The term describes the ventral germ layer of the trilaminar embryo, which will form the epithelium of the gut, including the prechordal plate, and many of their derivatives. These
possibly include prechordal mesenchyme but this may be of notochordal origin. Experimental studies suggest that the first cells ingressing through the primitive node give rise to notochord and embryonic endoderm,
which will form the roof of the secondary umbilical vesicle, displacing the cells of the hypoblast laterally into its walls. The qualifying adjective embryonic is necessary as long as terms such as primary endoderm
remain in use.
113 Lamina prechordalis The prechordal plate may not appear until Stage 7: certainly in Stage 8 it is “a highly developed mesendodermal mass [in which cells resemble either endoderm or mesenchyme] in
contact with the floor of the neural groove”. At stages 9 and 10, the plate is related to neuromere D1. Lateral growth at stages 9-11 gives rise to the bilateral premandibular condensations (Müller F, O'Rahilly R. The
prechordal plate, the rostral end of the notochord and nearby median features in staged human embryos. Cells Tissues Organs 2003;173:1-20). Prechordal mesenchyme does not become truly plate-like until Stage
9 and some would thus say that the prechordal plate appears in Stage 9.
114 Diverticulum allantoicum; Ductus allantoicus Several examples of “allantoic diverticula” have been reported in embryos of Stage 6. Nevertheless, “it is difficult to find a convincing example of an allanto-enteric
diverticulum at Stage 6” (O'Rahilly R, Müller F. Developmental stages in human embryos. Washington DC: Carnegie Institution of Washington; 1987).
116 Embryo postgastrulationis [St.6b ad 23] The pregastrulation and postgastrulation phases of the embryonic period and the fetal period are stages of prenatal development, each with its own distinctive
characteristics, particularly in respect of its responses to teratogens. In postgastrulation embryos the main events of organ formation occur. These entail delicate and complex tissue interactions that are particularly
susceptible to teratogens, which have dramatic effects upon morphology. Palate and lips, eyes, ears, brain, spinal cord and heart are all highly susceptible. Susceptibility diminishes as the main events of organ
formation are completed by Carnegie Stage 23.
117 Embryo cum processu notochordali [Gradus VII]; Embryo cum processu axiali Gradus VII]; Embryo cum chordomesoderma Gradus VII] Each embryo of Carnegie Stage 7 has a notochordal process
immediately rostral to its primitive node and streak. The embryos are generally about 0.4mm in diameter and about 19 days old.
118 Cardo chordoneuralis; Punctum chordoneurale cardinis Once the primitive node has started to form, gene expression centred on it becomes asymmetrical and the molecular basis for left-right asymmetry is
established. For a review of the first description by Hensen, see: Viebahn C. Hensen’s node. Genesis 2001: 29;96-103. See also: Müller F, O'Rahilly R. The primitive streak, the caudal eminence and related
structures in staged human embryos (Cells Tissues Organs; 177:2-20:2004); Charrier J, Teillet M, Lapointe F, Le Douarin N. Defining subregions of Hensen's node essential for caudalward movement, midline
development and cell survival (Development 1999; 126:4771-4783); Cambray, N, Wilson, V. Axial progenitors with extensive potency are localised to the mouse chordoneural hinge (Development 2002; 129:4855-
66).
119 Canalis neurentericus The neurenteric canal is a temporary passage between the amniotic cavity and the umbilical vesicle. It develops during Stage 8 and is constant (whether patent or not). It is most clearly
visible at Stage 9 and has almost disappeared at Stage 10 (Müller F, O’Rahilly R. The primitive streak, the caudal eminence and related structures in staged human embryos. Cells Tissues Organs 2004;177:2-20).
120 Embryo presomiticus [St.8] Embryos of Carnegie Stage 8 are late presomite embryos. They are generally 1-1.5mm in greatest length and about 23 days old. The term presomite embryo is sometimes applied
more generically to include also Stages 6-8 but this usage is not recommended. Originally, phases were not ascribed to Stage 8 (O'Rahilly R, Müller F. Developmental stages in human embryos. Washington DC:
Carnegie Institution of Washington; 1987). However, only advanced specimens show a neural groove: in the same specimens the floor of the notochordal process is breaking down and a notochordal plate is
present (O'Rahilly R, Müller F.The first appearance of the human nervous system at stage 8. Anat Embryol 1981;163:1-13). These distinct phases are recognised here by the use of the terms Presomite embryo
without neural groove [St.8a] and Presomite embryo with neural groove [St.8b].
121 Fovea primitive The term primitive pit is widely used but may be a misuse of the term primitive, which more usually refers to phylogeny rather than ontogeny. The term notochordal pit does not have this
anomaly and is more informative.
122 Ectoderma embryonicum The term describes the dorsal germ layer of the somite embryo, which will form the epithelium of the skin and nervous system and their derivatives. Experimental studies suggest
that, after obvious primitive streak activity ceases, epiblastic cells continue to ingress through the most caudal part of the primitive streak until the early somite stage. They form axial dense mesenchyme and thence
become the endoderm and mesoderm of more caudal parts in secondary body development. The cells remaining on the dorsal surface of the embryo thereafter constitute the embryonic ectoderm. The qualifying
adjective embryonic is necessary as long as terms such as primary ectoderm remain in use.
123 Somitomerum Somitomeres are paired whorls of mesenchymal cells that appear metamerically in paraxial mesoderm before the appearance of epithelial somites. Somitomeres form in strict craniocaudal
sequence beginning in the head where they subsequently contribute to head mesenchyme. Elsewhere, they condense, epithelialise and form somites. They have been found in all amniote embryos that have been
examined by stereo scanning electron microscopy (Jacobson AG. Somites and head mesoderm arise from somitomeres. In: Sanders EJ, Lash JW, Ordahl CP. Eds. The origin and fate of somites. Amsterdam: IOS
Press; 2001).
124 Zona junctionalis mesenchymalis The bar of mesenchyme where somatopleuric and splanchnopleuric mesenchymes meet and which separates the embryonic and extra-embryonic coeloms on each side of
the Stage 9 embryo. It breaks down, allowing them to communicate, in Stage 10.
126 Tuberculum caudale In Stage 20 the caudal tubercle (see above) is produced by vestiges of the nonvertebrated part of the caudal eminence or tail bud and may become cystic.
127 Mesenchyma intermedium; Mesoderma intermedium The tissue primarily responsible for the formation of the kidneys and internal genitalia and their ducts. Experimental studies suggest that intermediate
mesoderm in amniotes arises from the middle of the primitive streak distal to somite origin and proximal to lateral plate mesoderm origin. The term intermediate mesenchyme is thus more appropriate than the
commonly used intermediate mesoderm.
128 Cellulae antecedentes The term antecedent cell is used here solely in a generic sense and without any specific connotation. The term precursor is not used in a generic sense, to avoid confusion with the
specific term cellula precursoria, a synonym for the official term for stem cell, cellula staminalis.
129 Potestas totalis Totipotency is the capacity (a) to form all cell lineages, embryonic and extra-embryonic, and (b) to self-organise, i.e., to form the body axis and the basic body plan (e.g., zygote; blastomere).
130 Potestas pluralis Pluripotency is the capacity to form either all embryonic or all extra-embryonic cell lineages (e.g., cells of the inner cell mass or pluripotent stem cells).
131 Potestas multiplex Multipotency is the capacity to form multiple cell types of one cell lineage (e.g., cells of one germ layer; somatic adult stem cells).
132 Potestas una Unipotency is the capacity of adult stem cells to form only one cell type (e.g., haematopoietic stem cells; spermatogonia).
133 Formabilitas Plasticity is the ability of a specific stem cell population to transdifferentiate, i.e, to switch to a stem cell population with a different differentiation potential.
134 Cellula primordialis A primordial cell is totipotent; the zygote and its immediate progeny are primordial cells.
135 Cellula fundatoria Founder cells are capable of contributing to the establishment of one or more cell populations.
136 Cellula progenitalis; Cellula proprecursoria A prestem cell is capable of contributing to the establishment of one or more stem cell populations.
137 Cellula staminalis; Cellula precursoria A stem cell is a constituent of a population that is capable of maintaining its own size while exporting an appropriate output of progeny to one or more cell lineages. The
term Cellula staminalis was adopted by FIPATas the preferred term.
138 Cellulae multipotentes et unipotentes Cells are here usually listed according to both their derivation and their potential: an exception is the epidermal neural crest cell stem cell [eNCSC], which is derived from
epidermis but is capable of giving rise to neural crest cells.
139 Cellula staminalis germinalis An embryo of Stage 6b showed a marked concentration of glycogen in the extra-embryonic endoderm of the secondary yolk sac and some of the cells may be primordial germ
cells (Hertig AT, Adams EC, McKay DG, Rock J, Mulligan WJ, Menkin MF. A thirteen-day human ovum studied histochemically. Am J Obstet Gynecol 1958;76:1025-1043).
140 Factores crescentiae The factors listed here are only representative but all are known to be active in normal embryogenesis and specific congenital anomalies are known to be associated with disturbances of
them (TGF-α). Whether or not these criteria are appropriate and whether or not other growth factors should be included here is debatable. The number of growth factors and their families that have been identified
continues to increase as does knowledge of their activities.
142 Crista neuralis Cells of the primary neural crest separate from the neurosomatic ectodermal junction to give rise to the mesencephalic, rhombencephalic and spinal neural crest down to S1. Following
secondary neurulation, cells delaminate from the surface of the secondary neural tube and give rise to spinal neural crest beyond S1 (O'Rahilly R, Müller F. The development of the neural crest in the human. J Anat
2007;211:335-351). Neural crest is here divided according to the adjacent part of the brain. The term circumpharyngeal neural crest is not used as it describes a migration route to the pharyngeal region, the outflow
tract of the heart and great vessels and much of the gut-associated crest derivatives. Furthermore, it is said to be in the posterior rhombencephalic region but the crest for the first two pharyngeal arches is mainly
associated with rhombomeres 2 and 4.
143 Structurae cristae neuralis Neural crest tissue is dealt with under General histogenesis because of the wide range and distribution of its derivatives outside the nervous system. Groups of cells, which behave
in a similar manner but arise from some placodes and by delamination from the optic and otic vesicles are classified here as neural crest-like cells. Neural crest cells and neural crest-like cells meld seamlessly into
neural crest complexes and are no longer morphologically distinguishable (O'Rahilly R, Müller F. The development of the neural crest in the human. J Anat 2007;211:335-351). Here the term neural crest is used in
sensu stricto and the term neural crest complex recognizes the dual lineage of its component cells.
144 Complexus cristae neuralis nasalis The nasal neural crest complex develops from the epithelium of the nasal placodes in Stage 13 and migrates towards the telencephalon, reaching it in Stage 15, at which
stage complex-derived olfactory fibres enter the region of the future olfactory bulb.
145 Complexus cristae neuralis opticae The optic neural crest complex develops from the optic primordium in Stages 11 and 12 at the level of Diencephalon 1 and is the only forebrain-derived neural crest-like
tissue.
146 Cellula staminalis cornealis Corneal stem cells come from the corneoscleral junction.
147 Complexus cristae neuralis mesencephalicae The mesencephalic neural crest complex appears at Stage 9 and at Stage 11 spreads out towards the frontonasal region where it mingles with the optic neural
crest complex.
148 Crista neuralis isthmica Neural crest cells, seen in the roof of the isthmic rhombomere in Stage 13, appear to be destined more for the leptomeninges than for the mesencephalic nucleus of the trigeminal
nerve.
149 Complexus cristae neuralis trigeminalis At Stage 10 neural crest cells migrate mainly from future rhombomere 2 but with contributions from adjacent future rhombomeres and with neural crest-like cells from
the overlying ectoderm form the trigeminal neural crest complex.
150 Complexus cristae neuralis faciovestibulocochlearis At Stage 11 some cells from the otic vesicle, representing the primordial vestibular ganglion, attach to the facial neural crest. At Stage 14 afferent fibres to
the geniculate ganglion and efferent fibres from the vestibular ganglion distinguish between the two parts; at Stage 15 the smaller primordial cochlear ganglion cells appear and fibres are present at Stage 16.
151 Complexus cristae neuralis facialis At Stage 10 neural crest cells migrate mainly from rhombomere 4 but with contributions from adjacent rhombomeres and with neural crest-like cells from the overlying
ectoderm form the facial neural crest complex.
152 Ossiculae auditus (partim) Thompson et al. suggested that the mesenchyme of the auditory ossicles may also originate as a whole or in part from the neural crest. (Thompson H, Ohazama A, Sharpe PT,
Tucker AS. The origin of the stapes and relationship to the otic capsule and oval window. Dev Dyn 2012;241:1396-1404.)
154 Complexus cristae neuralis glossopharyngealis; Complexus cristae neuralis vagalis At Stage 10 neural crest cells migrate mainly from the roof of rhombomere 6 and probably from adjacent rhombomeres and
meld with neural crest-like cells from the overlying ectoderm to form part of a continuous glossopharyngeal/vagal neural crest complex. In Stage 13, the glossopharyngeal and vagal parts separate and each
develops superior and inferior ganglia, of which only the superior ganglia are believed to be derived from neural crest per se.
155 Crista neuralis cardiaca At Stage 12 some neural crest cells from rhombomeres 6 and 7, which are in continuity ventrally with the hypoglossal neural crest, proceed via pharyngeal arches towards the truncus
arteriosus; at Stage 13 they are joined by neural crest and neural crest-like cells and, continuing beyond the inferior glossopharyngeal and vagal ganglia, migrate into the 3rd and 4th pharyngeal arches; these
components are interpreted as human cardiac neural crest tissue.
156 Cartilagines laryngeae The Vagal neural crest complex does not contribute to all of the laryngeal cartilages.
157 Aa. arcuum pharyngeorum The terms pharyngeal arch artery / arteries are preferred to those of aortic arch/es to avoid confusion with the definitive aortic arch.
158 Crista neuralis nervi accessorii At Stage 12 neural crest cells from rhombomere 7 migrate and form the neural crest of the accessory nerve, which, by Stage 13, extends uninterruptedly between the vagal
neural crest and the spinal neural crest.
159 Crista neuralis hypoglossalis; Crista neuralis occipitalis In Stage 10 neural crest cells migrate from rhombomere 8 and spread between occipital somites where they mingle with myotomic cells to form the
hypoglossal cell cord; at Stage 12 the cord has reached the 1st pharyngeal arch; and by Stage 16 it has reached the lateral lingual swelling.
160 Crista neuralis spinalis The spinal leptomeninges, mesenchyme of the neck, trunk and lower limb dermis and adipocytes were formerly attributed to the spinal neural crest but are now known to be derived
from somites (Christ B, Huang R, Scaal M. Amniote somite derivatives. Dev Dyn 2007;236:2382-2396).The traditional distinction between trunk and lumbosacral spinal neural crest seems inappropriate because the
trunk lumbar and 1st sacral spinal neural crest all form in the same way, whereas the remaining sacral and coccygeal spinal neural crest, which form after the caudal neuropore closes at Stage 12, do so by direct
outgrowth from the secondary neural tube. The caudal limit of the spinal neural crest descends with each Stage, as does the formation of primordial spinal ganglia, which lie at somite 19 at Stage 13 and 33 at Stage
14.
161 Via migrationis dorsolateralis Dorsolateral migration from the spinal neural crest passes between the surface ectoderm and the dermatomyotome.
162 Via migrationis ventrolateralis Ventrolateral migration from the spinal neural crest passes between the dermatomyotome and the sclerotome.
163 Via migrationis ventromedialis Ventromedial migration from the spinal neural crest passes between the sclerotome and the neural tube.
164 Mesenchyma somatopleurale Together with ectoderm, somatopleuric mesenchyme makes up the body wall. The unqualified term somatopleure is not recommended because it is ambiguous, having been
used to mean either the whole thickness of the body wall or only its mesenchymal component
165 Mesenchyma splanchnopleurale Together with endoderm, the splanchnopleuric mesenchyme makes up the walls of the gut-related viscera. The unqualified term splanchnopleure is not recommended
because it is ambiguous, having been used to mean either the whole thickness of the visceral wall or only its mesenchymal component.
166 Cellula adipocytoprogenetrix Adipose tissue not only develops from mesenchyme from various sources but also from fat cell progenitors derived from granulocyte macrophage colony-forming units (GM-CFU).
168 Osteoclastus Osteoclasts form by the fusion of osteoclast progenitor cells (Bar-Shavit Z. The osteoclast: a multinucleated, hematopoietic-origin, bone-resorbing osteoimmune cell. J Cell Biochem
2007;102:1130–1139).
169 Canalis cartilagineus Cartilage canals first appear in the early fetus and by 28 weeks all the larger masses of cartilage are permeated by them. They contain blood vessels surrounded by loose cellular tissue
and provide the osteoblastic tissue for ossification when this later occurs (Haines RW. Cartilage canals. J Anat 1933;68:45-64).
170 Ossificatio perichondralis diaphysialis Adjectives derived from nouns such as diaphysis, epiphysis and hypophysis are, in a strict grammatical sense, probably best constructed using the suffix –alis giving
diaphysalis, epiphysalis, hypophysalis and symphysalis. However, for reasons of terminological precedence and consistency, the spellings of diaphysialis, epiphysialis, hypophysialis and symphysialis have been
here retained.
171 Anulus perichondralis This term describes the site of periosteal activity around the cartilaginous bud of a bone, and later the periosteal activity around the diaphysial cartilage.
172 Extensio gemmae osteogenicae The spread is that which takes place through an erosion canal into the cartilage model.
173 Osteonum primarium; Osteonum secundarium Primary osteons are directly deposited by the periosteum and not in a preceding resorption cavity. As a result, unlike secondary osteons, primary osteons are
not limited by resorption or reversal lines. Secondary osteons are deposited in a resorption cavity and are limited by resorption or reversal lines.