The structure of lymphatic capillaries
1
in lymph formation
LEE V. LEAK
The Ernest E. just Laboratory of Cellular Biology and the Department
Howard University, Washington, D.C. 20059
T he important function of the lym­
phatic system líes in the fact that nor­
mal blood capillaries are permeable
to the proteins that are capable of
disturbing the normal Starling pat­
tern of exchange between the blood
capillary wall and the surrounding
tissue space. This process, if left un­
checked, would lead to edema forma­
tion. The constant loss of diffusible
substances and plasma proteins from
the blood capillaries in the intersti­
tíum, if allowed to continue, would
not only deplete the blood circulatory
system of its plasma colloids, but
would also disrupt the balance of
forces responsible for the onlrol of
fluid movement and the exchange of
gases, fluids, and materials across
the blood vascular wall. Thus, the
lymphatic vascular y tem ub erves
th body ti sues by con·tant.J y rem v­
ing the exce int r tilia! Cluid and
plasma pr tein that are not remo ed
at th v nular limb of the blood vascu­
lar system. In this way the lymphati
system prcvents a build-up of Lhe
componem within th inter titium,
accompli hing the maintenance of
Aui.d horneo tasis fo.- the vari u
tissues of the body.
Although we now have sorne ideas
regarding th overall ontributi n of
the lymphati to the maintenance of
fluid homeo·ta ·i in ti ue through­
out the body, still to be resolved are
the exact mechanisms by which these
vessels accomplish these tasks and the
precise details of lymphatic ontogeny,
their ultrastructural anatomy, per­
meability, and organization in various
regions of the body.
A consistent interpretation for
many of the above-mentioned ques-
INTERSTITIAL FLUID PRESSURE ANO DYNAMICS OF LYMPH FORMATION
2
•
of Anatomy,
ABSTRACT
The lymphatic vascular system consists of endothelial lined vessels which begin as
blind-end tubes or saccules that are located within the connective tissue areas.
This system serves as a one-way drainage apparatus for the removal of diffusible
substances as well as plasma proteins that escape the blood capillaries. If permitted
to accumulate, these escaped cornponents would deplete the circulatory system of
its plasma colloids and disrupt the balance of forces responsible for the control of
fluid movement and the exchange of gases and fluids across the blood vascular
wall. The lymphatic capillaries are strategically placed and anatomically constructed
to permit a continuous and rapid removal of the transient interstitial fluids, plasma
proteins, and cells frorn the interstitium. Structurally the lymphatic capillaries con­
sist of a continuous endothelium that is extremely attenuated over major aspects of
its diameter, except in the perinuclear region which bulges into the lumen. These
vessels lack a continuous basal lamina and maintain a clase relationship with the ad­
joining interstitium by way of anchoring filaments. The adjacent cells are extensively
overlapped and lack adhesion devices in many areas. When electron-opaque tracers
are injected intravenously (i.e., horseradish peroxidase and ferritin), subsequent
electron microscopic examination of tissues reveals the presence of tracer particles
within the interstitium and the Iymphatic capillary lumen. These particles gain access
into the lyrnphatic capillaries via two major pathways: J) the intercellular clefts of
patent junctions and 2) plasmalemrnal vesicles (pinocytotic vesicles). Another salient
feature of the lymphatic endothelial • cell includes the presence of numerous cyto­
plasmic filarnents, which are similar in morphology to the actin filaments observed
in a variety of cell types. The ultrastructural features of the lymphatic capillaries
are discussed in relation to their role in the removal of interstitial Auids and
particulate matter, and in the formation of lymph.-Leak, L. V. The structure of
lyrnphatic capillaries in lymph formation. Federation Proc. 35: 1863-1871, 1976.
tions seems to depend in large meas­ p n ibl. for lymphatic capillary
ure on a more explicít understanding permeability wiU be onsid red, and
of the ultrastructural anatomy of the rol of the lymphatic in the dy­
the lymphatic vessels, their inter­ namics of the formatíon of lymph
cellular junctions, and the topo­ and its subsequent return to the blood
graphical relationship assumed by vascular system will also be discussed.
the capillary wall and its intirnate
association with the adjacent connec­
tive tissue compartments. In this 1 From the American Physiological Society
report, the structure and function of ymposium on hiten;li/ial Fluid PuSSILl"e and
Dynamics of Lympl, Fomuition pre ·ent d at
the lymphatic vascular system are Lhe 59th Annual Meeting of the Fcderation
presented and discussed in relation of American Societies for Experimental
to the topographical relationship • Biology. Achmtic Gity, NJ, April 16, 1975.
2 upportcd b grant AI10639 from the
between the lymphatic capillary wall
National Ln ·tilllte of Allergy and Infectious
and the adjacent connective tissue. Diseases and HL13901 from the National
In addition, tbe mechanisms - re- Heart and Lung Institute.
1863
 ORGANIZATION AND
UL TRASTRUCTURE OF THE
L YMPHATIC CAPILLARY
The earlier studies of Hudack and
McMaster (23) as well as a number of
other investigators demonstrated a
rich plexus of lymphatic capillaries
within the connective tissue of the
dermis (Fig. la, b ). Lymphatic capil­
laries could not always be recognized
Fi gu re l. a.) Demonstrates a whole mount preparation of lymphatics f rom 1he guinea pig's
ear, injected with colloidal carbon to label the lymphatic plexus (from Leak ancl Burke
(38)). b) Micrograph of Epon embeclded section from guinea pig's ear which ill11s1rates
lymphatic capill�rics (L) within che dermis (from Leak and Burke (37)).
1864
with certainty in routine histological
preparation, especially since they
were often collapsed ancl very diffi­
cult to distinguish from the connec­
tive tissue spaces or the clear areas
often produced by the techniques
used for obtaining sections from
paraffin preparations. However,
when the electron microscope was
first brought to bear on the organiza­
tion and structure of the lymphatic
FEDERATION PROCEEDINGS VOL. 35, NO. 8 • JUNE 1976
vessels, earlier workers (Palay and
Karlin (46), French, Florey and
Morris (20), Casley-Smith and Florey
(7), fraley and Weiss (19), Papp et al.
(47)) described the lymphatic capil­
lari s as being er imila1· lO blo d
apillaries x pt L11,H th 1 mphati
capillaries ar x rem -'ly i1T guiar in
shape and contain a poorly developed
basal lamina. In our studies of dermal
lymphatic capillaries (3, 33, 37)
specific criteria were established for
differentiating lymphatics from blood
capillaries at both light and electron
microscopic levels. The lymphatics
possess a wider and more irregular
lumen than the accompanying blood
capillaries; also, they are linee! by a
continuous endothelium that is
extremely attenuatecl over the major
aspects of its diameter except in the
perinuclear region. The basal lamina
is discontinuous and lacking in many
areas. A close topographical rela­
tionship is maintained with the ad­
joining interstitium by way of anchor­
ing filaments that are attachecl to the
endothelial wall and extend to the
surrounding connective tissue areas.
In addition, many of the adjacent
endothelial cells are extensively over­
lapped and lack adhesion devices.
Although the space between the ap­
posing cells may reach distances up to
severa! microns to allow passage of
large particles and cells, there are
specialized regions at various intervals
to form macula adherens and also
periodic macula occludens.
As indicated above, it is the lym­
phatic capillary that extends as a blind­
end tube or saccule into the inter­
stitium. The vessels are located near
the site of cellular metabolism and
also have a close topographical rela­
tionship with the blood capillaries
and smaller venules. Here this one­
way drainage system is strategically
placed and is anatomically con­
structed to permit a continuous and
rapid removal of transient interstitial
fluids, plasma proteins, and cells. By
using improved techniques to study
the lymphatic vascular system, a num­
ber of investigators have examined
lymphatic capillaries in a variety of
tissues and organs; these include the
lymphatics of the diaphragm (20),
dermal lymphatics (7, 19, 31, 32, 37),
the heart (41), the testis (15-17, 42),
the ovary (43), the spleen (27), the
ii1L tine (5 1.1 46, 47), and th
kidne and liver (25). WiLh im¡ rove 1
111 thod of ti· u pr s-rvation,
thic:k Ep 1 • ·Lion • exarnin I wiLh Lhe
Ji ·In mi ro cop reveal enain a -
p cLs <>f' lhe l y mphatic capillary struc-
1 u r llrnt usually escape r cogniLion
in the much thicker sections pre­
pared from paraffin sections (Fig. lb).
When observed with the electron
microscope, the lympbatic capillary
wall is shown to be lined by a con­
tinuous endothelium (Fig. 2a), which
is extremely attenuated over tbe
major aspects of its diameter except
in the perinuclear region. It has a
much wider ar,id a more irregular lu­
men than the accompanying blood L U M E N
capillary and lacks a definitive basal
lamina. The nucleus is located in the
tbicker region of the endothelium
and has a scalloped outline when the
vessel is collapsed; however, in the
dilated capillaries the nucleus takes on
a smooth to an elliptical contour.
The cytocentrum which contains a
pair of centrioles and a Golgi com plex
is located in the perinuclear region.
The Golgi consists of crescent shaped
!amellar membranous structures with
numerous vesicles closely associated
with its periphery (Fig. 3). Two
centrioles are occasionally observed in
close association with the Golgi ap­
paratus with one oriented per­
pendi uhr to the oLher. This arrange­
mem is reminiscem f that for cen­
trioles in other tissues (13).
There is a sparse endoplasmic re­
ticulum in the endothelial cells (Fig. L
3), which is mainly represented by
cisternae of the rough variety. In
addition, ribosomes are scattered Figure 2. a) Cross• eclion of lymplrnLit: capillary from the guin 'ª pig's <!ar. OH: Lh • irregular
throughout the cytoplasm in single outlinc oí LIP lymphatic capillary wall whicb al ·o onsis1s of exuerncly auenuated arcas
units and as polyribosomes (Fig. 3). l> yo11d 1.he perinuclear region (arrows). Thc nudeus ( ) l,ulge· into th lumen of 1h
Since large amounts of endoplasmic vessel. Note: thc ·y1oplasmi org;1nclles wiLhin Lh perinu I ar rcgion. fooc.:hondria (M)
reticulum of the rough variety are are al o listributed tJ1roughout 1h 1hin 1·im of Lhe ytoplasm; intercellular jun ·Lion ar·
usually characteristic of cells that as indioited (J) (from eak (34)). b) Thi· clectron mi rograph illu u, tes 11 e :Uta hmen1 of
have a high rate of protein synthesis an hori11g filarncm (af) ncar Lhc im rcellular junction (J) ;md thcir CXL n ion imo th con•
for extracellular discharge (4, 26, 45), nective tissue (CT) areas (from Leak and Burke (38)).
the small amount of endoplasmic
reticulum in the endothelial cells of
the lymphatic capillaries perhaps
indicates a slow or very limited turn­
over of protein for these cells. tions characteristic of mitochondria served throughout other areas of the
Mitochondria occur throughout in other cell types, i.e., cristae are cytoplasm.
the perinucl ar area and are also formed by enfoldings of the inner The dermal lymphatic capillary
randoml di Lributed throughout the membranes (Fig. 4a). There is a endothelial cells are distinguished by
attenuated 1·im of cytoplasm (Figs. small p pulaLi n f microtubules in numerous cytoplasmic filaments that
2b, 3, and 4a). They display the the cytoplasm, u ually in Lhe vicinity measure 40 to 60A in diameter
usual form and interna! configura- of ·entri 1 • and o a ionally ob- (Figures 4a, 4b, and 5). These fila-

INTERSTITIAL FLUID PRESSURE AND DYNAMICS OF LYMPH FORMATION 1865

 Eviclence that contraction of the
lymphatic capillary enclothelium is
clue to intrinsic cytoplasmic com­
ponents is suggested from the pres­
ence of cytoplasmic filaments in
lymphatic endothelial cells which
are morphologically similar to the
actin-like filaments in a variety of cell
types (48) ancl the demonstration of
anti-rabbit smooth muscle actin in
endothelial cells of lymphalic vessels
(10).

SPECIALIZATION OF THE
LYMPHATIC CAPILLARY WALL
The lymphatic capillary is dis­
tinguished by the specialization of
its wall, which permits a unique asso­
ciation and communication with the
adjacent connective tissue environ­
ment. The endothelial cell junc­
tions are extensively overlapping. In
addition, the apposing cell mem­
branes are loosely adherent and are,
therefore, free lo slide past each
other to form patent intercellular
junctions (Fig. 6a, b, e). This provides
an uninterruptecl channel between
the interstitium and the lymphatic
lumen which accommodates the
rapid passage of excess interstitial
fluids and plasma proteins.
The endothelial lined vessel is held
in close apposition to the surrounding
interstitium by way of anchoring fila­
ments (Figs. 2b, 4a, and 5). These
filaments appear to insert within a
densely staining substance on the
externa! surface of the endothelial
Fi gu re 3. The Golgi apparatus (G) is locatecl in the perinuclear region in aclclition to vesicles
cell membrane, and they extend into
associatecl with thc periphery of the Golgi. Large vesicles (V) containing an electron opaque
substance are also observecl in the perinuclear region. The encloplasmic reticulum (cr)
the adjacent connective tissue among
of the rough variety is sparse; howevcr, numcrous polyribosomcs (r) appear throughout collagen and elastic fibers (Figs. 2b, 5).
thc cytoplasm. Cytoplasmic filaments (el) are recognizecl as srnall clots when cut in cross­ They are in close proximity to the
sections; plasmalernrnal invaginations (*) occur along the connective tissue as well as along elastic fibers, are similar to the fila­
the lurninal front (v). Overlapping intercellular junctions are helcl in close apposition by ments within the mantle comprising
maculae adherens (arrow). Note the numerous anchoring filaments (af) along the ablurninal the elastic fibers, and may possibly
surface of the vessel. constitute a component of this extra­
cellular fiber. In examining the rela­
tionship between the lvmphatic wall
and the surrounding connective tis­
ments are of an undetermined length lymphatic endothelial cells of other sue, Drinker and Field (12) suggested
and are widely dispersed in various tissues (5, 31). that the lymphatics were "fixed" to the
regions of the cytoplasm. They are Although the rhythmic contraction connective tissue by fine strains of
observed in the peripheral cytoplasm observed in the collecting vessel reticulum which would produce dis­
adjacent to the plasmalemma and are (18, 22, 30, 52) can be attributed to tortion in the lymphatic wall as a result
also found in the perinuclear regions, the smooth muscle cells within its of muscular movement and, there­
often in bundles that run parallel to tunica media, the contraction ob­ fore, cause openings for the entrance
the long axis of the cell giving the served in the dermal lymphatic of fluids from the interstitium. Sub­
appearance of an electron-dense capillaries by cinephotographic sequently, Pullinger and Florey (49)
band in these regions of the endo­ methods cannot be ascribed to a demonstrated an intimate relation­
th lium. Num rou • cytoplasmic similar coat since the tunica media is ship between the lymphatic wall and
filaments have al o I een observ- ed in lacking in these vessels (Fig. 2a). the interstitium in normal and edema-

1866
FEDERATION PROCEEDINGS VOL. 35, NO. 8 • JUNE 1976
Figure 4. a) Section or lymphatic capillary wall which illustrates thc arrangement or rnitochondria (m), cytopbsmic filarnents (el), a11d plas111alcrn1md
vesicles (v) which contain injected colloidal ferritin. Polyribosomes (r) are also dispersed throughout the cytoplasm. Injected tracer particlcs
(arrows) appear ll'ithin the conncctive tissuc (CT) area as well as in the lymphatic lumen (L). /J) A portion of the lyrnphatic capillary wall illustrating
the nu111erous cytoplasmic filarnenls. Note the close association of anchorirrg filamenLs (at) to the lympliaLic capillary wall.

INTERSTITIAL FLUID PRESSURE AND DYNAMICS OF L YMPH FORMATION 1867

tous conditions. The anchoring fila­
ments, therefore, provide a struc­
tural basis for "fixing" the lymphatic
capillary wall to the surrounding inter­
sti Lium. This structural arrangement
provicles an increase of fluicls and
plasma proteins within the interstitium
that would presumably exert ec¡ual
pressure around the circumference of
the lymphatic vessel, thereby increas­
ing the total fluid pressure within
the interstitial space. There woulcl
also be an accompanying movement
of collagen bunclles, elastic fibers, and
other connective tissue components in
arder to accommodate the increased
fluid volume within the interstitium.
This would expand the interstitial
space ancl thus produce tension on
the collagen bundles and elastic fibrils
Fi gu re 5. This electron micrograph illustrates the appearance of anchoring (af) filaments
along the ablurninal surface of the lymphatic wall ancl their extension into the surrounding
connective tissue (CT) between collagen and elastic fibers. A portion of the nucleus (N)
as well as encloplasrnic reticulurn (er), mitochondria (m), polyribosomes (r), ancl cytoplasmic
filaments (cf) are also clemonstratecl (from Leak ancl Burke (39)).
1868
in which the anchoring filaments are
firmly embedded. Likewise, tension
would be placed or transferred to
the lymphatic capillary wall in which
the anchoring filaments insert.
Therefore, as collagen ancl elastic
fibers are separated by the increasecl
interstitial fluicls cluring both normal
ancl inflammatory conclitions, sites
along the lymphatic walls to which
anchoring filaments were attached
would also be pullecl along with the
collagen and connective tissue fibers.
This would result in a widening of
the lymphatic capillary lumen as the
extensively ancl loosely overlapping
adjacent endothelial cell margins
are separated to provide patent
intercellular junctions (Fig. 6c ).
It is of interest that the injection of
FEDERATION PROCEEDINGS VOL. 35, NO. 8 • JUNE 1976
hyaluronidase into edematous foot
pads of rats caused the lymphatics
to collapse (6), presumably as a re­
sult of a solubilization of connective
tissue ground substance (i.e., glycos­
aminoglycans) ancl a subsec¡uent clis­
ruption of the anchoring filaments.
This would reduce the stabilizing
or "fixing" effect of the anchoring
filaments on the lymphatic wall. The
extensive overlapping of the adjacent
endothelial cells would also serve as a
reserve supply of lymphatic wall
that can be immediately utilized for
rapid and uninterrupted expansion
for dilatation of the lymphatic capil­
lary lumen. This mechanism greatly
increases the overall capacity of the
lymphatic system in providing an ef­
fective one-way drainage system to
accommodate fluid removal from the
interstitial area. The extent of
lymphatic capillary dilatation ancl the
volume transfer of fluid would vary
greatly with the physiological state,
these conditions being increased
during the inflammatory response
(40).
The inner portian of the overlap­
ping endothelial cell junction appears
to open and close as a one-way flap­
valve system that serves to open
clefts of the intercellular junctions
when the interstitial fluid pressure is
higher than that of the lymphatic
intraluminal pressure (3). Con­
versely, when the intraluminal pres­
sure is ec¡ual to or higher than the
interstitial pressure, the one-way
flap-valve is closed. Recent studies of
Guyton and those of Scholander,
Knox, and Ott (see this symposium)
provide additional evidence for
the changes of interstitial fluid
pressures in both normal and edema­
tous conditions. Studies of Zweifach
(53) demonstrated that the intra­
luminal pressure in the mesenteric
lymphatic capillary was O to 2.5 cm
of water and that this intraluminal
pressure increased as the caliber of
vessels became larger and the thick­
ness of the muscular coat increased
around collecting vessels.
Another surface specialization of
the lymphatic capillary wall is the
presence of the plasmalemmal in­
vaginations. These appear on both
the connective tissue and luminal sur­
faces of the endothelial cells (Fig. 4a,
b). The functional significance of
these vesicles will be discussed on the
following pages.
 LYMPHATIC CAPILLARY phological and ex peri mental evi­ work of other investigators (5, 11),
PERMEABILITY dence that extracellular materials may also suggests a similar role for che
be internalized by endothelial cells as plasmalemmal vesicles in the lym­
The lymphatic capillary endothelium well as by a variety of other cell types phatic enclothelial cells.
is capable of rapiclly engulfing large after the in-pocketing of the plasma Although substances are Lrans­
111olecules ancl paniculate malter rnembrane to form vesicles (1, 8, 14, portecl from the connective tissue
from the adjoining interslilial area 28, 50, 51). Recent evidence obtainecl front into the lumen, sorne of the
(23, 24). A number of investigators in our laboratory (36), as well as the particles also aggregate to form large
have taken advantage of Lhis property
to ascertain the mechanisms involvecl
in the rnovernent of Auicls ancl large
particulate substances from the con­ Figure 6. a) An ex1ensivel)' overlapping intercellular junc1ion (J) is illustra1cd in this electron
nective tissue into the lymphatic ves­ rnicrograph. /¡) Illustrates thc appearancc of horseradish peroxiclase reac1ion prnclucl at
seis (7, 16, 37). 30 rnin, after an intravenous i1 1jcction of 1he tracer which appears in 1hc co1 1 11ec1ive
,vhen electron-opaque suspen­ tissue (CT), cleft of intcrcellular junctions (.J), and vesicles (V). e) A patcnl intcrcellular
sions of various sizes (80 Á to 0.08 junclion (j) in a lympha1ic capillary wall during the early period of 1he i 11 1lamrnatory
µ,m) ::tre interstitially injected with response is illusu·,ucd in this micrograph. In spite ol the increased intcrs1itial íluids ancl
microinjection techniques, their plasma proteins, the lymphatic capillar)' wall is held in close apposition lo thc surroundi11 g
connecLive tissuc (CT) components by lhe anchoring filamenls (af). Note the electron
subsequent movement from thc inter­
dense ílocculent material (*) within the patent intercellular cleft ancl the lumen (L) of
stitium into the lymphatic capillary Lhe lyrnphatic capillary (from Leak ancl Burke (39)).
can be observecl in the living animals
with c:inephotographic: methocls ancl
subsec¡uently at the light (Fig. la, b)
ancl electron microscopic levels (Figs.
5 and 7). WiLhin seconds after the
interstitial injection of trypan blue or
colloidal carbon, the dermal lym­
phatic capillaries are clearly cle­
lineated bv their content of the clye or
particles �s the tracer advanees very
rapidly toward thc larger collecting
vessels. Al severa! seconcls to 1 hour
following the interstitial injection of a
suspension of tracer substance into
dermal tissues, the tracer particles
are present in the interstitiurn and
the lumen of lymphatic capillaries,
as well as intracellular vesicles of
vanous sizes (Figs. 5 and 7). Inter­
stitially injected colloidal ferritin,
thorium, carbon, and latex spheres
gain entrance into lymphatics by
two morphological passageways,
1) the clefts of intercellular junctions
ancl 2) micropinocytotic vesicles.
The role of plasmalemmal vesicles
(micropinocytotic vesicles) in the
transport of substances across endo­
thelial cells was prnposecl in 1953 by
Palacle (44). He described these
vesicles in endothelial cells of bloocl
capillaries and suggestecl that ma­
terials perhaps pass from one side
of the cells to the other within such
structures. Subsequently, Bennett (2)
advancecl the hypothesis that mem­
brane flow and vesiculation may be
important transpon mechanisms
in carrying particles and molecules,
inclucling ions, within, into, ancl out
of the cell by way of vesicles. Since
these initial studies, numerous in­
vestigators have provicled both mor-

INTERSTITIAL FLUID PRESSURE ANO DYNAMICS OF LYMPH FORMATION 1869


Figure 7. This electron micrograph illustrates the appearance of the lymphatic capillary
and surrounding connective tissue after the interstitial injection of colloidal thorium. Note
the presence of tracer throughout the connective tissue (CT) and within the lymphatic
lumen (L).
hcterophagic vacuoles. This i • shown
by th ac umulati n of inert particles
uch as carb n and thorium dioxide
which may aggregate to form very
large vacuoles that reach several mi­
crons in diameter and are still filled
with tracer substances at 6 to 12
months after the injection. One also
observes whorled membranous struc­
tures along with electron-dense
bodies within these vacuoles.
The second and major passageway
for the movement of substances from
the interstitium into the lumen is
by way of clefts of intercellular junc­
tions (Figs. 6c and 7). This clearly
?�monstrates a direct entry of the
mJected suspensions into the lym­
phatic capillary lumen.
1870
In pite of the very a curate on­
trol of pressures that can be main­
tained when using microinjectioo
techniques, the possibilities still exist
of sorne trauma due to the develop­
ment of unphysiological pressures in
the immediate vicinity of the injec­
tion site. This hazard has been over­
come by using intravenous injections
of peroxidase following the proce­
dures of Graham and Karnovsky (21)
in which peroxidase is rendered visi­
ble for observation in the light and
electron microscopes.
After injecting peroxidase in the
tail veins of mice, dermal tissue of
the ear was fixed and processed at
various time intervals (2 minutes to
24 hours) for localization of the reac-
FEDERATION PROCEEDINGS VOL. 35, NO. 8 • JUNE 1976
tion product according to the meth­
ods of Graham and Karnovsky (21).
• early as 2 minutes after the inje ·­
ti n of peroxidase, the reaction prod­
uct wa observed in the blood capil­
laries, interstitium and clefts of inter­
cellular junctions, and within vesicles
and lumen of the lymphatic capil­
laries. The maximal intensity for the
rea tion product was reached by 30
minuce , as shown in Fig. 6b. Within
6 hours, there was a marked decrease
in intensity of the reaction product
throughout the interstitium; how­
ever, the intensity of the enzyme stain­
ing within ch lumen f lymphatic
capillarie wa still appre iable.
The mov ment of intravenou ly
injected peroxidase, like the inter­
stitially injected tracers, demonstrates
entrance into lymphatic capillaries
by way of both vesicles and the clefts
of intercellular junctions. The pass­
age of ub tances across th on­
nective tis u lymphatic interface is
summarized in diagrammatíc form in
Fig.
The appearance of an intra­
venousl injected protein within the
inrer titium and the lymphatic
capillary within 1 minute and a
marked decrease in the intensity of
Lhe reaction pr duct after l hour pro­
vides morpholocrical evidence for a
rapid and constam movement of thi
protein across the blood tonn ctive
ti· ue lymph interface by way of
interceUular juncti ns and ve i les
within the endothelium. [i"C
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INTERSTITIAL FLUID PRESSURE ANO DYNAMICS OF LYMPH FORMATION 1871