Vol 10, Issue 11, 2017
Online - 2455-3891
Print - 0974-2441
NEW INSIGHTS OF MAMMARY GLAND DURING DIFFERENT STAGES OF DEVELOPMENT
MANOJ KUMAR JENA
1
, ASHOK KUMAR MOHANTY
2
*
1
Department of Biotechnology, Lovely Professional University, Phagwara - 144 411, Punjab, India.
2
Animal Biotechnology Centre, National
Dairy Research Institute, Karnal - 132 001, Haryana, India. Email: ashokmohanty1@gmail.com
Received: 17 June 2017, Revised and Accepted: 20 July 2017
ABSTRACT
Mammary gland is a unique organ with its function of milk synthesis, secretion, and involution to prepare the gland for subsequent lactation. The
mammary epithelial cells proliferate, differentiate, undergo apoptosis, and tissue remodeling following a cyclic pathway in lactation – involution –
lactation cycle, thus fine tuning the molecular events through hormones, and regulatory molecules. Several studies are performed on the mammary
gland development, lactogenesis, and involution process in molecular details. The developmental stages of mammary gland are embryonic, pre-
pubertal, pubertal, pregnancy, lactation, and involution. Major developmental processes occur after puberty with hormones and growth factors
playing crucial role. The two major pathways such as Janus kinases-signal transducer and activator of transcription pathway and PI3K-Akt pathway
play a major role in maintaining the lactation. The involution process is a well-orchestrated event involving several signaling molecules and making
the gland ready for subsequent lactation. The review focuses on findings with molecular details of different stages of the mammary gland development
and signaling pathways involved in lactation–involution cycle. Deep insight into the developmental stages of mammary gland will pave the way to
understand mammary gland biology, apoptosis, oncogenesis, and it will help the researchers to use mammary gland as a model for research on
various aspects.
Keywords: Mammary gland, Lactation, Involution, Apoptosis, Development.
INTRODUCTION
Mammary gland (modified sebaceous gland) is exocrine in nature
and made up of branched network of ducts ending in alveoli [1]. This
gland is an excellent system to study the molecular events occurring
in proliferation and differentiation of cells as well as cancer cell
development (oncogenesis) [2]. This organ is unique in the sense; it can
involute and regain its shape in each pregnancy-lactation-involution
cycle. Mammary gland development mainly occurs after birth regulated
by peptide and steroid hormones [3]. A complex cross-talk exists among
various proteins expressed in different stages (pregnancy, lactation,
and involution) during the functional mammary gland development.
Intensive efforts have been made to unravel the molecular mechanisms
involved in lactogenesis, apoptosis of mammary epithelial cells
(MECs), and organ remodeling [1,4-7]. A number of previous studies at
transcriptome level have been performed on this aspect. For example,
cDNA library prepared from pooled sample of bovine mammary
tissues - with a number of ESTs, paved the way for functional genomics
studies of mammary gland [8]. Similarly, DNA microarray study
comparing gene expression profiles of lactating and non-lactating
bovine mammary tissue revealed many novel and interesting
differentially expressed genes such as cell cycle regulators, mediators
of apoptosis, and oncogenes [9]. Microarray analysis of transcriptional
changes during pregnancy cycle of mouse mammary gland with a focus
on lactation–involution transition revealed putative roles of death
receptors and immune mediators in post-lactational regression [10].
Involution is associated with an immune cascade and acute phase
response genes such as LBP, CD14, and signal transducer and activator
of transcription (STAT)3 [11]. The CCAAT/enhancer binding protein
delta is a crucial regulator of pro-apoptotic gene expression positioned
with STAT3 at upstream and several pro- and anti-apoptotic genes at
downstream level. A number of differentially expressed genes were
identified by microarray analysis to understand the molecular events
during the transition from late pregnancy to lactation in the mammary
gland of Holstein cows [12].
It is observed that milk composition varies with stage of lactation which
is specific to each species [13]. Milk is rich source of bioactive peptides
(derived from milk proteins) which have antioxidant and antibacterial
property [14]. Thus, it is important to understand the molecular events
occurring during lactation which will help to unravel more about the
milk components. This review focuses on the molecular details of
mammary gland development with latest findings on changes in the
mammary gland during pregnancy and lactation. This will help the
researchers to understand mammary gland biology and use this organ
as a model for different research area.
FUNCTIONAL ANATOMY OF MAMMARY GLAND
Mammary gland is a cutaneous gland such as sebaceous and sweat
glands, producing milk to feed the young offspring in female.
Histologically, it is a compound tubuloalveolar gland in the advanced
mammals originating from the ectoderm and evolved from apocrine
glands [15,16]. Bovine mammary gland is composed of two halves
separated by the median suspensory ligament which primarily supports
the udder along with the lateral suspensory ligament. Blood supply
is mainly through the external pudendal artery, and venous drainage
through the venous circle located at the base of the abdominal wall [16].
Mammary gland is made up of a branching network of ducts that end in
alveoli. Alveoli are the basic functional units of the milking mammary
gland and are grouped as lobules which connect to the collecting duct
system. Mammary gland is a complex secretory organ with number
of cell types: Epithelial cells (form the ductal network); adipocytes
(constitute the fat pad); vascular endothelial cells (form blood vessels);
stromal cells, including fibroblasts; and different immune cells [17]. The
epithelial cells are of two types: Luminal and basal. Luminal epithelia
form alveoli and duct system, whereas the basal one form myoepithelial
cells involved in milk ejection.
DEVELOPMENT OF MAMMARY GLAND
Hormones play significant role in mammary gland development through
distinct stages (Fig. 1). The mammary gland mainly consists of fat and
connective tissue before puberty, and there is moderate elongation of
epithelia into the fat pad which is not hormone-dependent. Ovarian
steroid hormones stimulate the extension, and branching of ducts after
© 2017 The Authors. Published by Innovare Academic Sciences Pvt Ltd. This is an open access article under the CC BY license (http://creativecommons.
org/licenses/by/4. 0/) DOI: http://dx.doi.org/10.22159/ajpcr.2017.v10i11.20801.
Review Article