Mamma:ry Glands

The Placenta and

22

I. INTRODUCTION. The placenta and mammary glands are nOI part ofthe female reproductive system; however, they are closely related to the reproductive function of the female reproductive system. A. Functions of the placenta. The fetus develops in a closed system; the placenta Is the interface between rnother and fetus. All material entering or leaving the fetal system passes through the placenta, The important functions of the placenta are: 1. Respiration
2. Nutrlemasslrnllatlon

3. Wasle elimination 4. Water, electrolyte, and immunoglobulin 5. Hormone synthesis

B. Functiolll§ of the mammary glands. Mammary glands are modified sweat glands. Their most important functlon is the production of milk. for the In·fanl. Milk has two important functions. transport

1. Milk is a. complete ulinsand

food forme Infant; it provides all ~he essential nutrients and minerals. substances, immunologic protection. II is nch in immunoglobwhich protect the infant against enteric infections.

2. Milk also provides the infant with essential

other bacteriostatic

II .. THE PLACENTA
A. Theio.rming

placenta. After ovum fertilization and cleavage, the compact mass of cells called the morulacavltates to form a. blastocoel. At this time, the embryo is called a blas1ocyst. 1. Trophoblast fonnation a, The blastocyst has a thin epithelial wall called the IJrophoblast and an inner cell mass (e.mbryoblast), which projects into the blastocoel. b. The trophoblast forms the chorion, a shell of tissue around the developing embryo, and subdivides into a cellular layer ca lied the cytotrophoblast, wh ich begins to proliferate rapidly. c. Cvtotropboblastlc cellular mitoses form new cells, wh ich fuse to form a syncytiall.ayer called the syncytiotfophoblast. (Figure 22-1). (1) The syncytiotrophoblast, which forms the outermost shell of the developing embryo, is an invasive syncytium that degrades matemalendometrial epilheUal cells and blood vessels. (2) As maternal blood vessels break down, maternal blood bathes the surface of the syncytiottophoblasl. 2. ChorionicviUi fonnation a. Cytotrophoblastk cells continue to divide and fuse to produce syncytiotrophoblast. They also differentiate into the mesenchymal cells that form blood vessels and connective tissue fjbrobla~rs .. b. Finger-like projections from the chorion and surface lnvaglnations called prima.ry villi begin
to

c. Primary villi become secondary

form. Primary yilli lack connective tissue and fetal blood vessels (Figure 22-2).
villi as they are i:nvaded by connective

tissue cores. 271

272

Histo/OBY and Cell Bi%gy

~~~3~~~;n::~--Endomet(ial ,epl1hellum ..llll:!-t+--- Endometrial gland

Cytotrtlphoblasl Figure 22·':. Human conceptus during implentatlon. Cytotrophoblast !;;ells divide and ruse 10 form the svncytiOlrophoblasl, an invasive struclUre Ihat faciHtalesconCep!U5 implantatlon. The endometrial epithelium has il.lrnO'it completely healed over Ihe lrnplanlalionsite. (Reprinled from J'ohnson KE: Human De'4!/opmeflfaf Anatomy. Baltimore, Williams & Wilkins, 1986, p 94.1 they are invaded by developing fetal blood vessels (Figure 22-.3), e. Many long villi develop on one side of lhe chorion, forming the chorion f.rondosum, which Is the precursor of the definitive placenta. The other side of the chorion has few, very short villi, which comprise thechorien laeve or smooth pari of the chorion.

d. Secondary villi become tertiary villi as

B. The definitive placenta 1. The chorionic plate and decidual

plate

a. As villi develop, the placenta develops two tissue layers. The chorionic 'pilate develops on the felal side or the placenta and the decidual plate (basal plate) develops on the maternal side of the placenta (Figure 22-4). b. Cytolrophobl'asliccell columns connect the two plates. Vl!Ji project from these columns into the intervillous space.

f,i&ure 2'2-2. Scanning electron micrograph of human placental choriontc villi (CV) auaehed to cytolro-

,,11!U5 biOpsy from a 'O-wf'ek human

'I)regna n ty .

phoblasrlc cell columns (CCq, This specimen was gathered by chorionic

Chapter 22

TIll.' Plecem« end Ml1mm,lry Ctsnds

273

surrounding a layer of cytotrophoblast (CTS),. The cure of the villus

consists 01 lela I connecri,ve tissue (en and felal blood vessels (Bill.

Figure 22-3. Llght micrograph of .1 tertiary villu~ from a firsl-Irimest(:!r human placenta. The syncy.liolrophoblaSI. lSTlJ) ls a syncytial layer

c. Maternal blood vesselstbranches of uteri ne artery) penetrate-the decidua I pi ate and enter the intervillous spaces. d. The human placenta is called a hemochodal placenta because maternal blood 'bathes the

syncytiotrophoblast.

2. The decidua
a. Decidual cells. Certain endometrial stromal fibroblastic cells undergo dramatic transforrnatlon from stellate or fusiform fibroblasts to glycogen-rich polygonal decidualceUs (Figure 22-5), Decidual cells form a barrier around the embryo and may help limit the invasiveness of the- syncytiotrophoblast. b. Decidual regions. The decidua has three regions.
MaternElI blood' vessels DeCIdual plata

Ir'lleNlllous &'\----+Chorlonlc

space plate Figure 224. Chorionic development as it nears the formation of a placenra. The rhononlc surface hili ,In area of exten)ivt' lIillous d!?IIt'IQPrnent (,nonon Irondosum; and <1 smoother area With fewer villi (chorion Iaew). The dec Id ua I pi ille and chorionic plate have formed and

cvrcrrepnomesuc call column

-Uterine

wall

are connected by cv'totrophoblaslic cell columns. CReponled from

[ohnson K£: Humiln Developmt'nla./

Analomv. Bailimore, Wilkins, 1988, p 97.)

Wilham~

&

274

His/clogy and Cell Biology

Figure 22·5. Ught micrograph of de· Cidual cells (DO. Tilesela'lle,polygonal: cells diffe.~enliate from stroma:
fibrobla5~(f).

The decidua basalis is the portion that becomes incorporated into the definitive placenta. II isan integr.al:pan of the placenta and is delivered at term. Thus, the placenta contains maternal and fetal components. (2.) The decidua capsularis 'is the portion that encapsulates Ihe implantation site. It expands as the embryo grows and then fuses with the decidua parietalls. obllteratlng the uterine lumen and forming the decidua vera. (3) The decidua parielalis is the portion that forms everywhe.re in the endometrial. wall except over the implantation site.
(1)

C. Functions of the placenta 1. Nut'rient transport. All fetal nutrients including glucose, fatty acids, amino acids, nucteotides, vitamins, and minerals are derived from the maternal blood supply and are transported across the placenta. 2. Electrolyte transport. Waler crosses the placenta by diffusion and pinocytosis. Salts lost in fetal urine are replaced by placental transport. Small ions enter the placenta by diffusion and pinocytosis mediated by small membrane-bound vesicles. 3. Gas transport. Oxygen bound to maternal hemoglobm drffuses across the placenta, enters fetal blood, and binds to fetal hemoglobin. Carbon dioxide bound to fetal hemoglobin diffuses across the placenta. enters the maternal blood, and binds to maternal hemoglobin. 4. Hormone synthesis a, The placenta produces farge amounts or progesterone and supplants the corpus luteurn as a source of progesterone after the first trimester. Due to this steroid biosynthesis function, the syncytIotrophoblast contains abundant smooth endoplasmic reticulum and many mitochondria, which arc essential for steroid biosynthesis and ATP production. b. The placenta. synthesizes large amounts of human chorionic gonadoliropin (heG). (1) hCG is a glycoprotein hormone that mainta ins the female reproductive system in a stale suItable for pregnancy. (2) hCG is synthesized on rough endoplasmic reticulum; glycosylation is completed in the Goigi apparatus. These organelles are weI'! developed in the syncytiotrophoblast.

Chapter 12

The Placenta and M,lmmiJry Gland.';

275
which

c. The placenta produces another polypeptide hormone called placent.al ladolen, stimulates the developmentcf Ihe mammary glands in preparation for lactatton.

5. Immunoglobulin transport a. Materna] immunoglobulin G (lgC) is transported across the placenta and enters fetal circulalion functionally inlact. Some of the vesicles in Ihe syneyl lotrophoblast are involved in IgG transport. b. Maternal IgG passively immunizes the fetus against the bacterial antigens that the fetus is exposed to at birth. c. After birth, maternal IgG is cataboljzederrd replaced by felallgG. which is synthesized de novo as the infant is exposed to bacterial antigens.

D. Cir,culationin the placenta

1. Materna.1 blood enters the intervillous space from branches of the uterine arteries. The oxygen In maternal blood diffuses through the walls or felal capillaries in the placenta into fetal blood. 2. fetal capillaries drain into venules, which empty into (he umbilical vein. The urnblllcal vein carries oxygen-rich blood through the umbihcal cord to (he ductus venosus, which bypasses the liver and empties into the inferlor vena cava of the fetus. 3. Oxygen-depleted blood I'e:luming from the fetal circulatory system flows Ihrough arterioles into Ihe umbilical arteries (branches of the internal iliac arteriesland then into fetal capillary beds in the placenta.

III. THE MAMMARY GlAN.DS

A. Structure. Mammary glands are compound tubuloalveolar glands. 1. Mammary glands have a complex duel system and numerous glandular acini. a.. Approximately 20 large ladiferous ducts empty into each nipple. lacli~erDus ducts branch several limes and then connect to lactiferous sinuses. b. Mammary gland lobules consist of many acini .. lobules are drollined by ducts that bra nch from the lactiferous sinuses.

2. lactiferous ducts are lined by strarifled squamous epithelium near the nipple and by simple cubctdaleplthehurn in the more distal portions of ducts. Acini are comprised of simple cuboidal
epithelial cells. 3. In the pre-pubertal and non-lactating mammary gland, glandular cells and ductal cells are quite similar. During lactalion,acinar cells differentlale into secretorycells thai secrete milk, which distends the acini, and cells lining many of the ducts become secretory (Figure 22·6).

4. During lactation, secretory cells become columnar. They have numerous apical microvilli, lvsosomes.
abundant rough endoplasmic reticulum, a well-developed Golg.i apparatus, fat globules, and

S. Myoepithelial: cells are intraepiLhelial cells tl.e., they reside on the apical side of the basement membrane) In acini and ducts that contain numerous contractile rnkrolllaments. These microfilaments contract to help express secreted milk from acini and ducts.

6. The secretion of milk occurs by .apocrine secretion (i.e., a small part of the apical portion of
each secretory cell is released with Ihe secretion product), B. Function. Milk production immunologic functions. is the most essential function of mammary glands, Milk has nutritive and

1. Nutritive

function protein constituent of milk.

b. Lactose is the predornlnant carbohydrale nutrient and osmotic elemenl in milk.

3.. Casein is the predominant

d. Milk also provides an infant with water, electrolytes, and vitamins, It is particularly rich in
calcium and phosphates. which are bound in lipid micelles and have an important role in the skeletal development of the infant

c. Milk is rich in a variety of lipids. The lipids are secreted from lipid droplets and form micelles in milk. Lipid micelles provide nutrition and help bind calcium and phosphate ions.

276

Histologyand Cell Biology

Figure 22-6. Light micrographs of non-lactating (top) and lactating (bol10m) mammary glands. The dis-tended acini of the lactating mammary gland contain milk (Ml.

2. Immunologic function

a. Milk is rich in IgG and secretory immunoglobulin A (slgA). slgA is a dimeric form of IgA that is protected from enteric proteolysis by a secretory component polypeptide. These immunoglobulins protect the infant from the enteric infections that can cause diarrhea, dehvdration, or malnulrition. b. Milk contains nonspecific antibacteriaJ substances such as lactoperoxidase and lysozyme. c. Milk contains maternal lymphocytes, which presumably have an immunologic function; however, this function is poorly understood.

Chapter 22

The Placenta end Mammal)' Gland::;

277

STUDY QUESTIONS
Directions: The groups of questions below consist of lenered choices followed by severa I numbered items. For each numbered item select the one lettered' choice with which it is most closely associated. Each lettered choice ma.y be used once, more than once, or not at all. QUestions

1-5

Questions 6--9
Match each function below with the rnost approprlatesvncvriouophoblastic organelle.

Match each description below with the most appropriate component of milk. (Al Immunoglobulin G (8) Secreloryimmunoglobulin (e) Casein (D) Lactose ( E) Lipid micelles 1. A dimeric immunoglobulin lhat prevents diarrhea and enteric infections in the newborn 2. A muItimolecular aggregate that bi;nds calcium and phosphate in milk 3. The predominant protein constituent of milk

(A) Rough endoplasmic reticulum

A (H) Smooth endoplasmic

reticulum

(el Goigi apparatus

(0) Membrane bound vesicles ( E) Mitochondria

G. Involved in both steroid biosynthesis and ATf' production

7. Involved inglycosylation of human chorionic gonadotropin (heG) and production o( secretory

vesicles
8. Site where polypeptide chains for hCGand human placental lactogen are assembled 9. Involved in pinocytosis and immunog.lobulin

4. Manufactured by maternal plasma cells; enters fetal blood, where it provides passive bacterial immunity for the infant 5. The major osmotic conslituent of milk

transport

278

HistDlogy and Cell Biology

Questions 10-14
Match each description below with the appropriate lettered structure in the micrograph.

10. Contains fetal capillaries and is the site of gas exchange in the placenta 11. A cytotrophoblastic cell column

12. Receives blood from a branch of the uterine artery and contains many maternal erythrocytes 13. Carries maternal oxygen to the fetus 14. Decidual cells

Ch"pter

22

The PIi)cenliJ and Mdmmary Glands

279

ANSWERS AND EXPLANATIONS

1-5. The answers are: 1-8, 2-E, 3-e, 4-A, 5 ·D. [II1Bl Immu nogtobu lin G IIgGl is a component of breast milk. It is also manufactured by maternal plasma cells and transferred intact across the placenta mto fetal circulation. IgG provides the fetus and newborn with passive immunity until fetal plasma cells become functlonal. Secretory immunoglobulin A (sigAl lsa dlrnerlc form of immunoglobulin A (lgA) associated wltha secretory component that prevents proteolysis in the gastroirnestinal Ifact of the infant. It is manufactured by maternal plasma cells in mammary gland connective tissues. ~lgA passes.into milk and coats enteric bacteria, preventing their attachment to epithelial cells in the gastrointestinal tract of the newborn. Bables lacking slgA are prone to enteric infections and diarrhea. Casein is the predominant protein constituent of milk and is rnanufactured on the endoplasmic reticulum of mammary gland secretory cells. Lactose, an lmportan! nutrient, is a disaccharide and the major osmotic component of milk Lipid micelles are an important nutrient and perform a transport function by binding calcium and phosphate. &-9. The answers are: 6-E, 7-C, 8-A, 9-0. [II C 4[ The syncytiotrophoblast contains a complex collection of organelles. which perform the same general (unctions that they perform in other cells in the body. The rough endoplasrrnc reticulum 15 involved In the synthesis or the polypepude chains of secretory proteins, such as human chorionic gonadotropin (hCG) and placental lactogen. In conjuncl ion with mitochondria, the smooth endoplasmic reticu lum has an importanf role In steroid

biosvnthesi s.
The Golgl apparatus completes the glycosylation of hCG, a g.lycoprolein hormone. Membrane·bound vesic les help transport substances across the syncytiotrophoblast in both d irections. For example, pinocytosis and endocytosis transport water and IgG in, and exocvtosis transports poly. peptide hormones out. Mitochondria are the source of All" for many syncytiotrophoblast synthesis and transport functions and are involved In steroid brosvruhesrs.

10-14. The answers are: 10·8, 11-A, 12-D, 13-0, 14--C III BI The dellrutlve placenta has two plates of tissue. The decidual plate is on the maternal side of the placenta, and the chorionlc plate is 011 the fetal side of the placenta, Cvtotrophoblasttc cell columns (AJ connect the decidual and chorionic plates. Gas exchange occurs at the surface of the villi IBJ that project (rom cvtotrophoblastlc cells into the intervillous space. Placental villi contain fet"r capillaries surrounded by connective tissues and, therefore, contain fetal erythrocytes. Oxygenated maternal blood from branches of the uteri ne arteries (fow~ through Ihe decidual plate into the intervillous space CD), where il bathes the villi ..Fetal capillaries carry fet.31 ervthrocvtes, which cafry maternal oxygen from the placenta to the fetus and carbon dioxide trom the fetus to the placenta. A layer of decidual cells (q surrounds the fetus and Cetalmembranes. This layer limits the invasiveness of the syncytiotrophoblast.