Normoblastic Maturation: Erythropoiesis

ERYTHROPOIESIS Estimates of time spent at each stage suggest that it takes
Normoblastic Maturation about one week for the BFU-E to mature to the CFU-E and
another week for the CFU-E to become a pronormoblast,
Terminology which is the first morphologically identifiable RBC precursor.
RBCs are formally called erythrocytes. The nucleated precursors in
the bone marrow are called erythroblasts. They also may be While at the CFU-E stage, the cell completes approximately
called normoblasts, which refers to developing nucleated cells three to five divisions before maturing further.1 As seen later,
(i.e., blasts) with normal appearance. This is in contrast to the it takes approximately another 6 to 7 days for the precursors
abnormal appearance of the developing nucleated cells in to become mature enough to enter the circulation, so
megaloblastic anemia, in which the erythroblasts are approximately 18 to 21 days are required to produce a
called megaloblasts because of their large size.
mature RBC from the BFU-E.
Three nomenclatures are used for naming the erythroid
precursors. Erythroid precursors
Normoblastic proliferation, similar to the proliferation of

1. erythroblast terminology is used primarily in
other cell lines, is a process encompassing replication (i.e.,
Europe.
division) to increase cell numbers and development from
immature to mature cell stages.
2. normoblastic terminology used more often in the
United States, it has the advantage of being
The earliest morphologically recognizable erythrocyte
descriptive of the appearance of the cells.
precursor, the pronormoblast, is derived via the BFU-E and
CFU-E from the pluripotential stem cells. The pronormoblast
3. rubriblast terminology because it parallels the
is able to divide, with each daughter cell maturing to the next
nomenclature used for granulocyte development.
stage of development, the basophilic normoblast. Each of
these cells can divide, with each of its daughter cells
maturing to the next stage, the polychromatic normoblast.
Each of these cells also can divide and mature. In the
erythrocyte cell line, there are typically three and
occasionally as many as five divisions2 with subsequent
nuclear and cytoplasmic maturation of the daughter cells, so
from a single pronormoblast, 8 to 32 mature RBCs usually
result.
Maturation process
Erythroid progenitors
The morphologically identifiable erythrocyte precursors develop
from two functionally identifiable progenitors, burst-forming
unit–erythroid (BFU-E) and colony-forming unit–erythroid (CFU-
E), both committed to the erythroid cell line.
4. Nucleoli disappear. Nucleoli represent areas where the
Criteria used in identification of the erythroid ribosomes are formed and are seen early in cell development
precursors as cells begin actively synthesizing proteins. As RBCs mature,
the nucleoli disappear, which precedes the ultimate cessation
of protein synthesis.
Morphologic identification of blood cells depends on a well-
stained peripheral blood film or bone marrow smear. In 5. The cytoplasm changes from blue to gray-blue to salmon
hematology, a modified Romanowsky stain, such as Wright or pink. Blueness or basophilia is due to acidic components that
Wright-Giemsa, is commonly used. The descriptions that attract the basic stain, such as methylene blue. The degree of
follow are based on the use of these types of stains. cytoplasmic basophilia correlates with the amount of
ribosomal RNA. These organelles decline over the life of the
The stage of maturation of any blood cell is determined by developing RBC, and the blueness fades. Pinkness
careful examination of the nucleus and the cytoplasm. The called eosinophilia or acidophilia is due to accumulation of
qualities of greatest importance in identification of RBCs are more basic components that attract the acid stain eosin.
the nuclear chromatin pattern (texture, density, Eosinophilia of erythrocyte cytoplasm correlates with the
homogeneity), nuclear diameter, nucleus:cytoplasm (N:C) accumulation of hemoglobin as the cell matures. Thus the cell
ratio, presence or absence of nucleoli, and cytoplasmic starts out being active in protein production on the ribosomes
color. that make the cytoplasm basophilic, transitions through a
period in which the red of hemoglobin begins to mix with that
blue, and ultimately ends with a thoroughly salmon pink color
Nucleus-to-Cytoplasm (N:C) Ratio when the ribosomes are gone and only hemoglobin remains.
The nucleus-to-cytoplasm (N:C) ratio is a morphologic

feature used to identify and stage red blood cell and white
blood cell precursors. The ratio is a visual estimate of what
area of the cell is occupied by the nucleus compared with the
cytoplasm. If the areas of each are approximately equal, the
N:C ratio is 1:1. Although not mathematically proper, it is
common for ratios other than 1:1 to be referred to as if they
were fractions. If the nucleus takes up less than 50% of the
area of the cell, the proportion of nucleus is lower, and the
ratio is lower (e.g., 1:5 or less than 1). If the nucleus takes up
more than 50% of the area of the cell, the ratio is higher
(e.g., 3:1 or 3). In the red blood cell line, the proportion of
nucleus shrinks as the cell matures and the cytoplasm
increases proportionately, although the overall cell diameter
grows smaller. In short, the N:C ratio decreases.
As RBCs mature, several general trends affect their appearance

graphically represents these trends.
1. The overall diameter of the cell decreases.
2. The diameter of the nucleus decreases more rapidly than

does the size of the cell. As a result, the N:C ratio also
decreases.
3. The nuclear chromatin pattern becomes coarser, clumped,

and condensed. The nuclear chromatin of RBCs is inherently
coarser than that of myeloid precursors. It becomes even
coarser and more clumped as the cell matures, developing a
raspberry-like appearance, in which the dark staining of the
chromatin is distinct from the almost white appearance of the
parachromatin. This chromatin/parachromatin distinction is
more dramatic than in other cell lines. Ultimately, the nucleus
becomes quite condensed, with no parachromatin evident at
all, and the nucleus is said to be pyknotic.

Normoblastic Maturation: Erythropoiesis

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ERYTHROPOIESIS Estimates of time spent at each stage suggest that it takes

Normoblastic proliferation, similar to the proliferation of

The nucleus-to-cytoplasm (N:C) ratio is a morphologic

As RBCs mature, several general trends affect their appearance

1. The overall diameter of the cell decreases.

2. The diameter of the nucleus decreases more rapidly than

3. The nuclear chromatin pattern becomes coarser, clumped,

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