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Theoretical Paper
Theoretical Paper
Morong Campus
COLLEGE OF SCIENCE
THEORETICAL PAPER
PRESENTED BY:
JANELL E. LONTOC
II – MB BS BIOLOGY MAJOR IN MICROBIOLOGY
PRESENTED TO:
EDMUND B. CAPUZ
PROFFESSOR
I. Proposed Title
The Evolution of Mammary Glands
II. Rationale
This theoretical paper aims to explain how mammary glands and lactation evolved.
III. Objective
The purpose of this study is to determine why mammary glands secrete milk and its significance
in all mammalian species. Which of these studies provides more evidence for the existence of
mammary glands will also be shown.
V. Cons Literature
Apendix B
(I simply provided an insight in 2 pages because only Charles Darwin's theory of the mammary
gland was disputed and proved by others that his theory was wrong.)
VIII. Recommendations
Based from the conclusion and studies gathered, the following recommendation are hereby
suggested:
1. Further study on how mammary gland evolved and evolution of lactation.
2. More article explaining the idea of lactation in mammals.
3. More study focusing on the distinction of mammary glands.
IX. Notes
https://edisciplinas.usp.br/pluginfile.php/5663978/mod_resource/content/1/Oftendal.pdf
https://pdf.sciencedirectassets.com.pdf
https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2688910/
cambridge.org/core/journals/nutrition-research-reviews/article/evolution-of-lactation-
nutrition-v-protection-with-special-reference-to-five-mammalian-
species/9A518FD5D8D1BC4B711E34441939A394
https://www.ncbi.nlm.nih.gov/pmc/articles/PMC8515151/
X. Curriculum Vitae
Appendix C
Appendix A
Appendix B
Appendix C
In humans and other mammals, a mammary gland is an exocrine gland that produces milk to
feed young children. Mammals are named after the Latin word mamma, which means "breast." The
mammary glands are placed in organs such as primates' breasts (for example, humans and chimps),
ruminants' udders (for example, cows, goats, sheep, and deer), and other species' dugs (for example,
dogs and cats). Lactorrhea, or the occasional production of milk by the glands, can occur in any animal,
but lactation, or the production of enough milk for nursing, happens only in phenotypic females who
have gestated within the last few months or years in most mammals. It is guided by the hormonal
supervision of sex steroids. Male lactation is possible in a few mammalian species. Male lactation is
only possible in humans under certain conditions. Mammals are classified into three categories:
prototherians, metatherians, and eutherians. Both males and females have functional mammary glands
in prototherians, however their mammary glands lack nipples. These mammary glands are repurposed
sebaceous glands. Only females have functional mammary glands in metatherians and eutherians.
Breasts or udders are the names given to their mammary glands. Each mammary gland has its own
nipple in the case of breasts (e.g., human mammary glands). Pairs of mammary glands form a single
mass in udders, with more than one nipple (or teat) dangling from it. Cows and buffalo, for example,
have one udder with four teats, whereas sheep and goats have two teats extending from the udder.
Hormones control the development of mammary glands, the synthesis of milk, and the ejection
of milk to suckling infants. Mammary glands in eutherian mammals are typically found as paired
structures ranging in number from 2 to 18, and can be found on the thorax (human, elephant, bat),
along the entire ventral thorax and abdomen (sow, rabbit), in the inguinal region (horse, ruminants),
along the abdomen (whale), or even dorsally (whale) (nutria or coypu, a South American rodent). These
glands appear to be modified sweat glands, which are likewise unique to mammals. Internally, the
structure is very homogeneous, with supporting stromal cells and a glandular epithelium organized into
clusters of minute, sac-like structures known as alveoli. This glandular epithelium is in charge of the
production of milk. The alveoli are connected by ducts and other duct-derived enlargements for milk
storage. There are also modified epithelial cells with muscle-like myofilaments running parallel to the
cells' long axis. These cells, known as myoepithelial cells, are capable of contracting and ejecting milk
from the alveoli into the duct system and out of the gland in the nipple region.
Lactation appears to be a primitive reproductive trait that predates the evolution of mammals.
Olav Oftedal has proposed a compelling argument for the evolution of the mammary gland and
breastfeeding. The characteristics of modern mammals were gradually acquired through synapsid
ancestors' radiations, and the mammary gland is thought to have developed from apocrine-like glands
connected with hair follicles. According to Oftedal, these glands evolved from supplying moisture and
antimicrobials to parchment-shelled eggs to supplying nourishment for progeny. Evidence from the past
suggests that some therapsids and mammaliaformes, which lived during the Triassic period more than
200 million years ago, secreted a nutrient-rich milklike secretion. Lactation can be divided into two main
stages or phases that are controlled by distinct endocrine pathways. The first stage is lactogenesis, or
milk secretion. Pituitary PRL (or placental CS), growth factors, and glucocorticoids are the primary
regulators of this process. Lactogenesis is also induced by GH in primates. Lactogenesis is the process
by which milk fat, protein, and sugar, often lactose, are synthesized. Lactose synthesis is ultimately
dependent on protein synthesis; that is, lactose synthetase, the enzyme responsible for lactose
synthesis, must be activated. Lactose synthetase is composed of two protein units, one of which is
lactalbumin, which is also found in milk. Lactose, fat, and milk protein (mostly casein) are released into
the alveolar lumen. Osmosis allows water and other water-soluble compounds to enter the lumen,
resulting in a watery liquid known as milk. Milk contains a variety of hormones, such as hypothalamic
peptides, pituitary. Particularly intriguing is the impact that the disaccharide lactose (Galactose 1-4
Glucose) played in the development of lactation. Almost all foods include lactose milks, either in their
free form or at the reducing end of oligosaccharides (with the exception of a few marine animals' milks).
Its production in mammary glands is reliant upon the presence of a 1,4-galactosyl transferase and a
mammary-specific protein (-lactalbumin), the two work together to create the heterodimer lactose.
synthetase. Evidently, the enzyme lysozyme, which is widely distributed and breaks down glycosidic
bonds in bacteria, is where lactalbumin gets its starting material and was probably present in the
secretions of cell walls old glands that resemble apocrine glands hormones, growth factors, steroids,
However, despite what molecular data implies, Messer and Urashima found it difficult to
understand why -lactalbumin would have developed more than 100 million years before the appearance
of late Triassic "mammals" (or mammaliaforms). If early synapsid eggs benefited from gland secretions
that contained oligosaccharides, this problem would no longer exist. When the secretions began to
supply nutrition in addition to water and protecting chemicals, the mammary evolution was significantly
redirected. This is a unique usage of skin secretions that may have started during the egg-incubation
phase since it would improve egg uptake and the digestion of tiny carbs liquid intake. The lack of a
cemented eggshell in synapsid eggs indicates that the calcium needed for embryonic development
must have been placed in the egg yolk, as in squamates with eggs that have paper shells. However, it
appears that the calcium being provided hardly meets demands, leaving newly hatched upon hatching,
squamates require a calcium source in their diet. Contrast this with the seeming abundance of calcium
that is available to birds, turtles, and crocodiles with eggs made of calcium. Ionic calcium for eggs may
have been given by glandular secretions. At least in certain taxa, squamates' retained eggs may absorb
Caseins are tertiary, highly hydrated proteins found in mammalian milk. They contain clusters of
phosphoserine and phosphothreonine residues that bind amorphous calcium phosphate, allowing milk
to contain higher levels of soluble calcium than is possible in ordinary solutions. As a result of its highly
mobile structure, casein resembles a denatured protein in its native state in many ways, and its
allergenic activity is much less affected by thermal treatments than that of many globular proteins.
Casein linear epitopes are available for IgE binding both before and after thermal treatment. Short-term
boiling of milk lowers IgE binding to caseins only to a limited extent. Caseins most likely evolved as a
source of amino acids, phosphate, and calcium for hatchlings rather than embryos, because casein
micelles are too big to pass through eggshell pores. Although the origins of casein genes are unknown,
S1-, S2-, and -casein genes may represent duplication and divergence of a single starting gene, and
the -casein gene may be derived from the -fibrinogen gene. Because mammaliaforms' restricted tooth
replacement implies significant calcium intake for bone formation prior to independent feeding,
The initial milk produced by monotremes and some marsupials is quite diluted (about 10-12%
dry matter) and includes low quantities of both lipids and lactose, as well as monosaccharides, are
simple sugars. Initial expression of the glycosyltransferases needed for oligosaccharide synthesis is
lacking. This early secretion differs greatly from the high oligosaccharide and high lipid milks that are
released in later lactation stages and may be the most similar to a "primitive" milk to be found in extant
mammals. Before the end of the Triassic, milk must have developed into something more or less similar
to its contemporary composition, and it must have been essential to the reproductive success of both
early mammaliaforms and mammals, as well as mature cynodonts. Before cynodonts could get smaller
and more endothermic, two prerequisite steps that came before the emergence of tiny mammaliaforms
in the late Triassic and early Jurassic, lactation had to emerge as a dominating trait.
The idea that breast secretion has an ancient beginning and a long evolutionary history is gaining
traction, especially in light of supporting molecular evidence. It is now conceivable to develop a more
thorough scenario for how the produced fluid came to resemble what we currently call milk in structure
and function. Milk's evolutionary beginnings appear to be found in the secretions of primitive apocrine-
like glands in the skin of early synapsids or even taxa living prior to the split of synapsids from
sauropsids around 310 million years ago. Over the course of several radiations of early synapsids and
the following lineages leading to mammals, this secretory fluid and the glands that made it got more
sophisticated, the volumes produced increased, and the level of reliance by hatchlings and developing
young increased. Endothermic incubation of eggs with parchment-like shells appears to have required
the existence of a glandular skin secretion. In terms of metabolism, locomotor capacity, structural
adaptations to nutrition, and reproduction, the radiations of basic synapsids, therapsids, cynodonts, and
of species (e.g., cows, sheep, goats, water buffaloes, yaks, camels, horses, and asses) with relatively
dilute milks containing modest levels of fat, caseins, and whey proteins, and relatively high levels of
lactose as dairy animals. Perhaps such species were chosen because their milks were similar to human
milk in that they were dilute, or because their dilute milks collected in large storage cisterns between
bouts of suckling. However, it is only now that we are realizing how long the extraordinary secretory
product that is the backbone of the dairy industry has existed before mammals came on the planet.
Cattle, buffaloes, goats, sheep, and camels produce almost all of the world's milk. Yaks, horses,
reindeer, and donkeys are other less frequent milk animals. Each species' prevalence and importance
vary greatly among regions and countries. The main factors that influence the dairy species retained
are feed, water, and climate. Market demand, dietary customs, and the socioeconomic features of
individual homes are all factors that may influence the prevalence of a dairy species (e.g., poorer
families tend to rely more on small ruminants). Although cattle are raised in a variety of habitats, other
dairy species allow dairying to be practiced in unfavourable situations that cannot often support any
other sort of agricultural output. Sheep provide milk in semi-arid places around the Mediterranean, goats
in African regions with poor soils, horses in Central Asian steppes, camels in desert lands, buffaloes in
wet tropical regions, and yaks in high mountainous areas such as the Tibetan Plateau. Milk-producing
animals are frequently raised in subsistence and smallholder systems in developing nations. These
animals are typically multipurpose and develop and produce under challenging conditions such as low
inputs, minimal management, and severe surroundings. They are well acclimated to local environments,
Around 270 million dairy cows are cared for by millions of farmers worldwide who produce milk.
The extent to which milk production affects the environment depends on the methods used by dairy
farmers and feed producers. Climate change is a result of greenhouse gas emissions from dairy cows
and their excrement. Local water supplies might be harmed by improper treatment of manure and
fertilizers. Additionally, unsustainable dairy farming and feed production can result in the destruction of
forests, wetlands, and other ecologically significant habitats like grasslands. According to WWF, all
dairy products will be produced as sustainably as possible and sold on a global scale. WWF wants to
change the milk production industry by working to involve dairy farmers, co-ops, businesses, and others
amniotes had no skull windows, such as species that had dual temporal fenestrae and were related to
crocodilians, and birds that are still alive today. Thus, the divide between the Synapsida and the other
taxa (referred to as the Sauropsida collectively) is older than 310 million years. To put this in
perspective, the earliest evidence of bone, a feature of vertebrates, dates from the late Cambrian, which
is just 200 million years earlier. It is crucial to understand that mammals and reptiles diverged from the
earliest amniotes, not vice versa. We shouldn't anticipate living reptiles to closely resemble their
mammalian ancestors in anatomy, physiology, or behavior. The synapsids have undergone numerous
radiations and extinctions in the 300 million years since their first appearance, forming the dominating
fauna in the Permian and Triassic until the dinosaurs overtook them in the late Triassic. Around 225
mya, the first mammal-like or mammaliaform synapsids appeared. True mammals first appeared during
the Jurassic and Cretaceous, when dinosaurs were still prevalent, and only fully developed in the
Tertiary, when the dinosaurs had vanished End-Cretaceous ecological crisis led to extinction. At some
time during this protracted period of separation, lactation likely developed in response to selecting
pressures that promoted greater parental commitment in the offspring. It is crucial to understand that
during the extensive evolutionary history, numerous modifications took place from the earliest synapsid
to the earliest mammal. The best studies focus on those that are visible in the fossil record. Numerous
of these reflect modifications in nutrition (particularly predatory specialization), mobility, and energy
requirements. These changes didn't happen all at once. Synapsids have an exceptionally good fossil
record, which shows repeated radiations of taxa incorporating an increasing number of mammalian
traits. The emergence of these traits through time in the fossils that gave rise to mammals is frequently
cited as a prime illustration of gradual, continuous evolution. This shows connected progression, in
which a variety of traits progressively change in an interdependent manner. It is likely that such a
significant advancement as lactation evolved gradually rather than by a saltational jump, and that its
development into a period of intensive nutrient transfer was linked to the evolution of other traits that
now distinguish mammals, such as an accelerated metabolic rate, high aerobic capacity, rapid nutrient
Over and above the selection benefits afforded by milk's nutritional and antibacterial
characteristics, the mammary glands and the lactation process have played an important role in the
evolutionary success of mammals. Lactation allows for maternal effects on development and the final
phenotype of the child to occur in addition to those that occur during pregnancy or via behavioral
interactions. Lactation entails significant metabolic costs, which emerge as parent-offspring conflict,
and special physiological adaptations have developed to adjust milk supply to demand by the young.
The publication explicitly states that the mammary gland evolved from a compelling proposal for the
evolution of the mammary gland and lactation has been proposed. Olav Oftedal contributed the
information. The characteristics of modern mammals evolved gradually through radiations of synapsid
progenitors, and the mammary gland is thought to have originated from apocrine-like glands connected
with hair follicles. According to Oftedal, these glands evolved from supplying moisture and
suggests that therapsids and mammaliaformes produced a nutrient-rich milklike secretion during the
Triassic epoch more than 200 million years ago. These nutrients had to be supplied by milk due to the
shrinkage of the amniotic yolk. It's possible that some of these changes started occurring rather early
ironbinding proteins, and certain oligosaccharides into glandular secretions may have been favored by
natural selection since the feeding of fluids to eggs would have produced a favorable environment for
microbial attack. Early mammary secretions may have had a similar purpose to some sauropsid eggs'
substantial albumen percentage, which serves to protect the egg from microbial and fungal attack while
Milk may be excellent for bone growth, but it also had a significant impact on mammalian
evolution. A new genetic study backs up the assumption that nursing evolved before mammals
abandoned their reptile relatives' egg-laying habits. And milk may have aided in the biological
modifications that enabled the move to live birth. Mammals appeared approximately 200 million years
ago. Most of them eventually replaced their shell-bound eggs with a placenta to nurture the child in
gestation. They also started raising their children with milk. Monotremes, such as the platypus, are living
exceptions since they produce milk but still lay eggs. Milk secretion is a property shared by all
mammalian species, whether large or little, gregarious or solitary, arctic or tropical. Carolus Linnaeus,
the great taxonomist, chose the name Mammalia in 1758 to unite terrestrial 'quadrupeds' with dolphins
and whales to represent the fact that females of both mammae and mammary glands, are found in
clusters. These creatures are grouped together. Lactation is a tremendously intricate evolutionary
process that appears to be quite old origin. The intricate signaling interplay essential for mammary
tissue development, ductal branching, and proliferation is only now being unraveled, and the functional
importance and patterns of expression of thousands of mammary genes are being investigated.
Secreted milk contains unique proteins (a-, b-, and k-caseins, b-lactoglobulin, a-lactalbumin, whey
acidic protein (WAP)), membrane-enclosed lipid droplets, and sugars (lactose, milk oligosaccharides)
Milk also carries an important set of innate immune system components that help the neonate
transition from the relatively sterile environment of the mother's uterus to a postnatal environment
functional diet that contains many growth factors, hormones, and chemicals that act as metabolic
signals to the newborn, as well as enzymes and cofactors of comparable complexity to those found in
the cytosol of the offspring. Furthermore, human milk contains approximately 130 different
oligosaccharides and over 600 different fatty acid molecules. Milk also improves offspring survival by
maturation in the newborn Milk appears to meet the offspring's immediate and long-term demands in
this aspect. These requirements can vary greatly per species. Suckling and nurturing between mother
(dam) and offspring also has behavioral and 'psychological' features that increase neonate survival.
This is a unique feature of lactation regardless of the chemical and physical properties of milk Vorbach
and others highly conserved inflammatory response to tissue damage and infection, according to al,
may have aided in the evolution of lactation. As a result, scientists determined that the evolutionary
sequence was designed to give protection first, and then nutritional components evolved to nourish
Oftedal postulated that the evolutionary antecedent to milk was released by hypertrophied skin
glands connected with hair follicles and the skin. Natural selection favored secretion because it
protected the synapsids' parchment-shelled eggs from desiccation and microbial attack. Furthermore,
this secretion gradually became nutrient rich, which coincided with a progressive decrease in egg size,
suggesting a possible dual purpose providing some nutritional transfer during egg incubation and, later,
an enteral source of nutrients for hatchlings. Lactation may have developed to avoid parchment shell
egg desiccation by spreading a thin coating of fluid from glands associated with hair follicles over the
egg. Oftedal also proposed that nipples were incompatible with distributing fluid over the egg and that
projecting hairs interfered with suckling when viviparity emerged. Thus, nipples may have developed in
lineages that produced live-born neonates. Neonatal suckle is enabled by adaptations such as a
secondary hard-palate and jaw modifications, and nipples give a simpler attachment site than a breast
patch.
The frequency of suckling varies greatly between species, and behavior patterns have developed
to suit the maternal physiology. Furthermore, the frequency of suckling in is highly variable in women.
Similarly, lower-producing beef cows are suckled five times per day, whereas high-producing dairy cows
are customarily milked twice per day without notable adverse effects. As a result, suckling differs widely
between and within species. Nonetheless, the change in suckling frequency implies that there must be
intriguing control mechanisms governing the rate of milk synthesis so that it may adapt to variations in
child appetite and suckling intervals. The similarities in the morphology of feeding structures, both
newborn (oral anatomy) and maternal (mammary gland architecture), reflect a common feeding
process. Most mammals' mammary glands have a conspicuous nipple or teat to which the young
latches to remove milk. Indeed, nipples and suckling appear to be a later development in breastfeeding,
possibly coinciding with the greater nutritional role. When milking, nipples may have conferred an
advantage production increased because they prevent milk leakage and aid in milk removal by
Reproductive success in animals is based on, first, synchronizing birth with the onset of copious
milk production, and, second, maintaining lactation until the young is able to build a competent host
defense system and has developed enough to be sufficiently fed an adult diet. Furthermore, despite
function conservation, the energy demands of lactation are significant, and the nursing mouse produces
its body weight in lipids during the 20-day lactation period. Even in women, when newborn growth is
relatively slow, the lactating breast takes roughly 30% of resting energy.
I've suggested that milk evolved from an egg supplement to a food source for hatchlings.
However, it likely performed both functions in egg-laying synapsids, and it may still do so in living
monotremes. Future studies that show the transfer of mammary secretions from the monotreme
mammary areolar patch to eggs, as well as the uptake of water and/or nutrients by these eggs, will
support the evolutionary scenario envisioned here and further our understanding of the evolution of
mammary glands.
A first objection to Darwin's theory of evolution by natural selection was answered by his
explanation of how lactation may have originated. The issue was that the evolution of lactation was
impractical because a neonate could not gain from ingesting a mother's accidental secretion in order to
In response, Darwin proposed that mammary glands descended from cutaneous glands that
were housed within the brood pouches used by some fish and other marine species to store their eggs,
giving the eggs nutrition and a better chance of surviving. Because it has persevered through this and
other difficulties 200 years after Darwin's birth, the theory of evolution by natural selection continues to
be a pillar of biology. In response to Mivart's critiques, Darwin dedicated the majority of a chapter in the
sixth 1872 edition of On the Origin of Species to them of mammary development. Darwin saw that eggs
are hatched and raised in a brood pouch in seahorses (Hippocampus sp.), and he believed the
young the cutaneous secretions in this pouch might provide nourishment. He questioned, "Is it not
possible that the young might have been similarly nourished? Is it not possible that mammals are
In this instance, the individuals that released a fluid were the most nutrient-dense in order to
mimic the properties of milk would eventually have produced more well-nourished offspring than would
those who secreted a poorer fluid, and as a result, the cutaneous glands, which are the mammary
glands' counterparts, would have been enhanced or made more effective, and the glands over a specific
area of the sack should have developed to a higher degree. As we can observe in the Ornithorhynchus
[platypus], at the base of the mammalian series, they would have first produced a breast without a
nipple. Despite the fact that Darwin had observed a platypus while serving as a naturalist on the Beagle
expedition in Australia in 1836, the idea that this species deposited shelled eggs was regarded as
implausible because no other animal was known to do so. The presence of huge paired oviducts and
enormous ovaries implied ovi-viviparity, or the retention of eggs in the uterus until they hatch and then
emerge as young, similar to many squamates (lizards and snakes). It became obvious that monotreme
reproduction encompassed both egg-incubation and lactation, being a curious blend of avian and
mammalian features, when W. H. Caldwell verified in 1884 that the monotremes (platypus and
echidnas) were actually egg-laying. Darwin's theory of the origin of breastfeeding was in tatters if
lactation could evolve in an egg-laying animal and if this animal was similar to a platypus in that it lacked
a pouch. Although Darwin's argument of the evolution of the mammary gland did not hold up under
Personal Background
Age: 22
Sex: Female
Date of Birth: April 06, 2000
Place of Birth: Cardona, Rizal
Religion: Roman Catholic
Civil Status: Single
Nationality: Filipino
Father’s name: Nolan F. Lontoc
Mother’s Name: Evangeline E. Lontoc
Educational background
Tertiary: University of Rizal System 2021 - Present
Morong, Rizal
Major: Biology Major in Microbiology
Our Lady of Fatima University 2018 - 2020
Antipolo, Rizal
Major: Medical Laboratory Science
Secondary: Renaissance School of Science and Technology 2012 – 2018
Morong, Rizal (Junior and Senior High)
Elementary: Looc Elementary School 2006 - 2012