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Bone marrow
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Bone marrow

A simplified illustration of cells in bone marrow

Details
Latin

Medulla ossium
Identifiers

MeSH D001853 (https://www.nlm.nih.gov/cgi/mesh/2011/MB_cgi?mode=&term=Bone+Marrow)

TA

A13.1.01.001
(http://www.unifr.ch/ifaa/Public/EntryPage/TA98%20Tree/Entity%20TA98%20EN/13.1.01.001%20Entity%20TA98%20EN.htm)

FMA

9608 (http://fme.biostr.washington.edu/FME/index.jsp?fmaid=9608)
Anatomical terminology

Bone marrow is the flexible tissue in the interior of bones. In humans, red blood cells are produced by cores of bone marrow
in the heads of long bones in a process known as hematopoiesis. On average, bone marrow constitutes 4% of the total body
mass of humans in an adult weighing 65 kilograms (143lb), bone marrow typically accounts for approximately 2.6 kilograms
(5.7lb). The hematopoietic component of bone marrow produces approximately 500 billion blood cells per day, which use the
bone marrow vasculature as a conduit to the body's systemic circulation.[1] Bone marrow is also a key component of the
lymphatic system, producing the lymphocytes that support the body's immune system.[2]
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Bone marrow transplants can be conducted to treat severe diseases of the bone marrow, including certain forms of cancer
such as leukemia. Additionally, bone marrow stem cells have been successfully transformed into functional neural cells,[3] and
can also potentially be used to treat illnesses such as inflammatory bowel disease.[4]

Contents
1 Structure
1.1 Types
1.2 Stroma
1.3 Cellular components
2 Function
2.1 Mesenchymal stem cells
2.2 Bone marrow barrier
2.3 Lymphatic role
2.4 Compartmentalization
3 Society and culture
4 Clinical significance
4.1 Disease
4.2 Examination
4.3 Donation and transplantation
4.3.1 Harvesting
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5 Fossil record
6 See also
7 References
8 Further reading

Structure
Types
The two types of bone marrow are "red marrow" (Latin: medulla ossium rubra), which consists
mainly of hematopoietic tissue, and "yellow marrow" (Latin: medulla ossium flava), which is
mainly made up of fat cells. Red blood cells, platelets, and most white blood cells arise in red
marrow. Both types of bone marrow contain numerous blood vessels and capillaries. At birth,
all bone marrow is red. With age, more and more of it is converted to the yellow type only
around half of adult bone marrow is red. Red marrow is found mainly in the flat bones, such as
the pelvis, sternum, cranium, ribs, vertebrae and scapulae, and in the cancellous ("spongy")
material at the epiphyseal ends of long bones such as the femur and humerus. Yellow marrow
is found in the medullary cavity, the hollow interior of the middle portion of long bones. In
cases of severe blood loss, the body can convert yellow marrow back to red marrow to
increase blood cell production.

Stroma

A femoral head with a cortex

of cortical bone and medulla of
trabecular bone. Both red bone
marrow and a central focus of
yellow bone marrow are visible.

The stroma of the bone marrow is all tissue not directly involved in the marrow's primary function of hematopoiesis. Yellow

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The stroma of the bone marrow is all tissue not directly involved in the marrow's primary function of hematopoiesis. Yellow
bone marrow makes up the majority of bone marrow stroma, in addition to smaller concentrations of stromal cells located in
the red bone marrow. Though not as active as parenchymal red marrow, stroma is indirectly involved in hematopoiesis, since it
provides the hematopoietic microenvironment that facilitates hematopoiesis by the parenchymal cells. For instance, they
generate colony stimulating factors, which have a significant effect on hematopoiesis. Cell types that constitute the bone
marrow stroma include:
fibroblasts (reticular connective tissue)
macrophages, which contribute especially to red blood cell production, as they deliver iron for hemoglobin production.
adipocytes
osteoblasts
osteoclasts
endothelial cells, which form the sinusoids. These derive from endothelial stem cells, which are also present in the bone
marrow.[5]

Cellular components
Cellular constitution of the red bone marrow parenchyma[6]
Group

Cell type

Average Reference
fraction
range

Myeloblasts

0.9%

0.2-1.5

Promyelocytes

3.3%

2.1-4.1

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Myelopoietic
cells

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Neutrophilic myelocytes

12.7%

8.2-15.7

Eosinophilic myelocytes

0.8%

0.2-1.3

Neutrophilic metamyelocytes

15.9%

9.6-24.6

Eosinophilic metamyelocytes

1.2%

0.4-2.2

Neutrophilic band cells

12.4%

9.5-15.3

Eosinophilic band cells

0.9%

0.2-2.4

Segmented neutrophils

7.4%

6.0-12.0

Segmented eosinophils

0.5%

0.0-1.3

Segmented basophils and mast cells 0.1%

0.0-0.2

Pronormoblasts

0.6%

0.2-1.3

1.4%

0.5-2.4

21.6%

17.9-29.2

Orthochromatic normoblast

2.0%

0.4-4.6

Megakaryocytes

< 0.1%

0.0-0.4

Plasma cells

1.3%

0.4-3.9

Reticular cells

0.3%

0.0-0.9

Lymphocytes

16.2%

11.1-23.2

Monocytes

0.3%

0.0-0.8

Erythropoietic Basophilic normoblasts

cells
Polychromatic normoblasts

Other cell
types

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Hematopoietic precursor
cells: promyelocyte in the
center, two
metamyelocytes next to it
and band cells from a bone
marrow aspirate.

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In addition, the bone marrow contains hematopoietic stem cells, which give rise to the three classes of blood cells that are
found in the circulation: white blood cells (leukocytes), red blood cells (erythrocytes), and platelets (thrombocytes).[5]

Function
Mesenchymal stem cells
The bone marrow stroma contains mesenchymal stem cells (MSCs),[5] also known as marrow stromal cells. These are
multipotent stem cells that can differentiate into a variety of cell types. MSCs have been shown to differentiate, in vitro or in
vivo, into osteoblasts, chondrocytes, myocytes, adipocytes and beta-pancreatic islets cells.

Bone marrow barrier

The blood vessels of the bone marrow constitute a barrier, inhibiting immature blood cells from leaving the marrow. Only
mature blood cells contain the membrane proteins, such as aquaporin and glycophorin, that are required to attach to and pass
the blood vessel endothelium.[7] Hematopoietic stem cells may also cross the bone marrow barrier, and may thus be harvested
from blood.

Lymphatic role
The red bone marrow is a key element of the lymphatic system, being one of the primary lymphoid organs that generate
lymphocytes from immature hematopoietic progenitor cells.[2] The bone marrow and thymus constitute the primary lymphoid
tissues involved in the production and early selection of lymphocytes. Furthermore, bone marrow performs a valve-like function
to prevent the backflow of lymphatic fluid in the lymphatic system.

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Compartmentalization
Biological compartmentalization is evident within the bone marrow, in that certain cell types tend to aggregate in specific
areas. For instance, erythrocytes, macrophages, and their precursors tend to gather around blood vessels, while granulocytes
gather at the borders of the bone marrow.[5]

Society and culture

Animal bone marrow has been used in cuisine worldwide for millennia.

Clinical significance
Disease
The normal bone marrow architecture can be damaged or displaced by aplastic anemia, malignancies such as multiple myeloma,
or infections such as tuberculosis, leading to a decrease in the production of blood cells and blood platelets. The bone marrow
can also be affected by various forms of leukemia, which attacks its hematologic progenitor cells.[8] Furthermore, exposure to
radiation or chemotherapy will kill many of the rapidly dividing cells of the bone marrow, and will therefore result in a
depressed immune system. Many of the symptoms of radiation poisoning are due to damage sustained by the bone marrow cells.
To diagnose diseases involving the bone marrow, a bone marrow aspiration is sometimes performed. This typically involves using
a hollow needle to acquire a sample of red bone marrow from the crest of the ilium under general or local anesthesia.[9]

Examination
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Bone marrow examination is the pathologic analysis of samples of bone marrow obtained via biopsy and
bone marrow aspiration. Bone marrow examination is used in the diagnosis of a number of conditions,
including leukemia, multiple myeloma, anemia, and pancytopenia. The bone marrow produces the cellular
elements of the blood, including platelets, red blood cells and white blood cells. While much information
can be gleaned by testing the blood itself (drawn from a vein by phlebotomy), it is sometimes necessary to
examine the source of the blood cells in the bone marrow to obtain more information on hematopoiesis
this is the role of bone marrow aspiration and biopsy.
A Wright'sstained bone
marrow aspirate
smear from a
patient with
leukemia.

The ratio between myeloid series and erythroid cells is relevant to bone marrow function, and also to
diseases of the bone marrow and peripheral blood, such as leukemia and anemia. The normal myeloid-toerythroid ratio is around 3:1 this ratio may increase in myelogenous leukemias, decrease in polycythemias,
and reverse in cases of thalassemia.[10]

Donation and transplantation

In a bone marrow transplant, hematopoietic stem cells are removed from a person and infused into
another person (allogenic) or into the same person at a later time (autologous). If the donor and
recipient are compatible, these infused cells will then travel to the bone marrow and initiate blood
cell production. Transplantation from one person to another is conducted for the treatment of
severe bone marrow diseases, such as congenital defects, autoimmune diseases or malignancies. The
patient's own marrow is first killed off with drugs or radiation, and then the new stem cells are
introduced. Before radiation therapy or chemotherapy in cases of cancer, some of the patient's
hematopoietic stem cells are sometimes harvested and later infused back when the therapy is
finished to restore the immune system.[11]

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A bone marrow harvest in

progress.

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Bone marrow stem cells can be induced to become neural cells to treat neurological illnesses,[3] and can also potentially be used
for the treatment of other illnesses, such as inflammatory bowel disease.[4] In 2013, following a clinical trial, scientists
proposed that bone marrow transplantation could be used to treat HIV in conjunction with antiretroviral drugs[12][13] however,
it was later found that HIV remained in the bodies of the test subjects.[14]
Harvesting
The stem cells are typically harvested directly from the red marrow in the iliac crest, often under general anesthesia. The
procedure is minimally invasive and does not require stitches afterwards. Depending on the donor's health and reaction to the
procedure, the actual harvesting can be an outpatient procedure, or can require 12 days of recovery in the hospital.[15]
Another option is to administer certain drugs that stimulate the release of stem cells from the bone marrow into circulating
blood.[16] An intravenous catheter is inserted into the donor's arm, and the stem cells are then filtered out of the blood. This
procedure is similar to that used in blood or platelet donation. In adults, bone marrow may also be taken from the sternum,
while the tibia is often used when taking samples from infants.[9] In newborns, stem cells may be retrieved from the umbilical
cord.[17]

Fossil record
The earliest fossilised evidence of bone marrow was discovered in 2014 in Eusthenopteron, a lobe-finned fish which lived
during the Devonian period approximately 370 million years ago.[18] Scientists from Uppsala University and the European
Synchrotron Radiation Facility used X-ray synchrotron microtomography to study the fossilised interior of the skeleton's
humerus, finding organised tubular structures akin to modern vertebrate bone marrow.[18] Eusthenopteron is closely related to
the early tetrapods, which ultimately evolved into the land-dwelling mammals and lizards of the present day.[18]
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See also
National Marrow Donor Program, a nonprofit organization that operates a registry of
volunteer hematopoietic cell donors and umbilical cord blood units in the United States
Gift of Life Bone Marrow Foundation, an American bone marrow transplantation registry

References

Bone marrow may have first

evolved in Eusthenopteron, a
species of prehistoric fish
with close links to early
tetrapods.

1. Vunjak-Novakovic, G. Tandon, N. Godier, A. Maidhof, R. Marsano, A. Martens, T. P. Radisic, M.

(2010). "Challenges in Cardiac Tissue Engineering". Tissue Engineering Part B: Reviews 16 (2): 169.
doi:10.1089/ten.teb.2009.0352 (https://dx.doi.org/10.1089%2Ften.teb.2009.0352).
2. The Lymphatic System (http://allonhealth.com/health-news/par-lymphatic-system.htm).
Allonhealth.com. Retrieved 5 December 2011.
3. "Antibody Transforms Stem Cells Directly Into Brain Cells"
(http://www.sciencedaily.com/releases/2013/04/130422154756.htm). Science Daily. 22 April 2013.
Retrieved 24 April 2013.
4. "Research Supports Promise of Cell Therapy for Bowel Disease" (http://www.wakehealth.edu/NewsReleases/2013/Research_Supports_Promise_of_Cell_Therapy_for_Bowel_Disease.htm). Wake
Forest Baptist Medical Center. 28 February 2013. Retrieved 5 March 2013.
5. Raphael Rubin and David S. Strayer (2007). Rubin's Pathology: Clinicopathologic Foundations of
Medicine (http://books.google.co.uk/books/about/Rubin_s_Pathology.html?
id=kD9VZ267wDEC&redir_esc=y). Lippincott Williams & Wilkins. p.90. ISBN0-7817-9516-8.

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6. Appendix A:IV
(http://www.msd.com.mx/secure/ebooks/WintrobesClinicalHematology/sid4266054.html) in
Wintrobe's clinical hematology (9th edition). Philadelphia: Lea & Febiger (1993).
7. "The Red Cell Membrane: structure and pathologies" (http://www.acbd.monash.org/docs/red-cellmembrane.pdf) (PDF). Australian Centre for Blood Diseases/Monash University. Retrieved
24 January 2015.
8. "Human acute myeloid leukemia is organized as a hierarchy that originates from a primitive
hematopoietic cell" (http://www.nature.com/nm/journal/v3/n7/abs/nm0797-730.html). Nature.
1997. Retrieved 9 November 2012.
9. "Bone Marrow Aspiration and Biopsy"
(http://www.labtestsonline.org.uk/understanding/analytes/bone-marrow/tab/sample). Lab Tests
Online UK. Retrieved 16 February 2013.
10. "Definition: 'M:E Ratio' " (http://www.medilexicon.com/medicaldictionary.php?t=75938). Stedman's
Medical Dictionary via MediLexicon.com. 2006. Retrieved 20 December 2012.
11. "Bone marrow transplantation" (http://www.uptodate.com/contents/bone-marrow-transplantationstem-cell-transplantation-beyond-the-basics). UpToDate.com. Retrieved 12 April 2014.
12. "Bone marrow 'frees men of HIV drugs' " (http://www.bbc.co.uk/news/health-23132561). BBC. 3
July 2013. Retrieved 3 July 2013.
13. "Stem-Cell Transplants Erase HIV In Two Men" (http://www.popsci.com/science/article/201307/stem-cell-transplants-erase-hiv-two-men). PopSci. 3 July 2013. Retrieved 3 July 2013.
14. "HIV Returns in Two Men Thought 'Cured' by Bone Marrow Transplants"
(http://rhrealitycheck.org/article/2013/12/10/hiv-returns-in-two-men-thought-cured-by-bonemarrow-transplants/). RH Reality Check. 10 December 2013. Retrieved 10 December 2013.

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15. National Marrow Donor Program Donor Guide

(http://www.marrow.org/DONOR/When_You_re_Asked_to_Donate_fo/index.html). Marrow.org.
Retrieved 5 November 2012.
16. Bone marrow donation: What to expect when you donate (http://www.mayoclinic.com/health/bonemarrow/CA00047). Mayo Clinic. Retrieved 16 February 2013.
17. McGuckin, C. P. Forraz, N. Baradez, M. -O. Navran, S. Zhao, J. Urban, R. Tilton, R. Denner, L.
(2005). "Production of stem cells with embryonic characteristics from human umbilical cord blood".
Cell Proliferation 38 (4): 245255. doi:10.1111/j.1365-2184.2005.00346.x
(https://dx.doi.org/10.1111%2Fj.1365-2184.2005.00346.x). PMID16098183
(https://www.ncbi.nlm.nih.gov/pubmed/16098183).
18. Sanchez S., Tafforeau P. and Ahlberg P. E. (2014) "The humerus of Eusthenopteron: a puzzling
organization presaging the establishment of tetrapod limb bone marrow"
(http://rspb.royalsocietypublishing.org/content/281/1782/20140299). Proceedings of the Royal
Society B: Biological Sciences 281 (1782): 20140299. doi:10.1098/rspb.2014.0299
(https://dx.doi.org/10.1098%2Frspb.2014.0299)

Further reading
Nature Bone Marrow Transplantation (http://www.nature.com/bmt/index.html) (Nature
Publishing Group) specialist scientific journal with articles on bone marrow biology and clinical
uses.
Cooper, B (2011). "The origins of bone marrow as the seedbed of our blood: from antiquity to

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Bone marrow.

the time of Osler" (http://baylorhealth.edu/proceedings/24_2/24_2_Cooper.pdf). Baylor University Medical Center

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Proceedings 24 (2): 1158. PMC3069519 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3069519). PMID21566758

(https://www.ncbi.nlm.nih.gov/pubmed/21566758).
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Categories: Bone marrow Skeletal system Hematopoiesis Endocrine system Lymphatic system Stem cells
Lymphoid organ
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