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Therefore, we performed a detailed evaluation of the parameters of
the contractility of aged mesenteric lymphatic vessels in vivo and char-
acterized lymph flow in the aged mesenteric lymphatic network [100].
Results were compared with the same characteristics of lymphatic
contractility and flow in adult mesenteric lymphatic vessels. We per-
formed these evaluations under control conditions and after a nitric
oxide synthases blockade of 100-μM
L-NAME. This allowed us to
make important conclusions about the comparative roles of the
NO-dependent regulatory mechanisms, which control the lym-
phatic contractility and lymph flow in vivo in the adult and aged
body. Importantly, we performed, for the first time, the direct
detailed comparison of the characteristics of the lymphatic contrac-
tility in vivo and contractile characteristics of the isolated lymphatic
vessels exteriorized from the same location for both adult and aged
MLV under both control conditions and after NOS synthases
blockade.
On the other hand, a careful analytical comparison of the influ-
ences of the NO-synthase blockade in the aged MLV in vivo and
in isolated vessel-based studies allowed us to determine the impor-
tant differences (discussed later) between functioning of the lym-
phatic vessels under isolated conditions and functioning of the
same vessels under the additional influence of the aged tissue
microenvironment.
In a follow-up study, we evaluatedthe aging-associatedchanges in
the functional role of histamine as an EDRF in aged MLVs [105].We
measured and analyzed parameters of lymphatic contractility in iso-
lated and pressurized rat mesenteric lymphatic vessels (MLVs)
obtained from 9- to 24-month-old Fischer-344 rats under control con-
ditions and after pharmacological NO blockade by Nω-nitro-
L-
arginine methyl ester hydrochloride (
L-NAME, 100 μM) or/and
blockade of histamine production by α-methyl-
DL-histidine dihy-
drochloride (α-MHD, 10 μM). We also quantitatively compared
results of immunohistochemical labeling of the histamine-producing
enzyme histidine decarboxylase (HDC) in adult and aged MLVs.
This study [105] demonstrated that aging introduces significant
changesin wall shear stress-dependentmodulationof lymphatic con-
tractility related to the increased functional role of histamine as an
EDRF in aged MLVs and simultaneous effective loss of the NO as
a molecular player in such regulatory reactions. These findings
84 4. Hydrodynamic regulation of lymphatic vessel transport function
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correlate with the increased expression of HDC, the histamine-
producing enzyme, in aged MLVs. At the same time, we confirmed
that transmural pressure-/stretch-dependent regulatory reactions
of contracting MLVs are not affected by intralymphatic-borne hista-
mine, neither in adult nor in aged MLVs. Such is not true for effects
of histamine delivered from sources external to lymphatic vessel on
the lymphatic contractility. In particular, data from the literature sug-
gest that histamine is a potent dose-dependent modulator of lym-
phatic contractility [83,90,91,106–111]. At lower concentrations,
histamine acts synergistically with the lymphatic-relaxing effects of
NO, but at higher concentrations, histamine acts predominantly
antagonistically to NO and stimulates lymphatic contractility
[83,90,91,106–116]. Such findings illustrate the clear necessity of fur-
ther in-depth discovery-driven research efforts to establish compli-
cated interrelations between histamine-producing lymphatic vessels
and histamine-producing cellular elements of perilymphatic tissue
compartments.
The findings of the current study arose from the experiments,
which clearly illustrate the continued necessity of describing the
many unknown features of lymphatic vessels to gain insight into
the origin and pathogenesis of diseases of the lymphatic system,
for example, lymphedema. As we mentioned earlier [84], histamine,
as a known chemoattractant of mast cells [117], may be responsible
for increased density of mast cells observed near MLVs [77]. In light
of our current findings, an observed further increase of mast
cell density in aged perilymphatic mesenteric tissues (compared
with adult tissues) [77] directly correlates with the increased expres-
sion of HDC in aged MLVs demonstrated in this study. Increased
levels of histamine in aged MLVs, with corresponding increased
diffusion of this molecule out of lymphatic vessels, are very likely
important for increased recruitment and potentially for chronic
basal activation of perilymphatic mast cells in the elderly [77,118].
Lastly, recent data demonstrate that the aging-associated basal acti-
vation of perilymphatic mast cells maintains chronic histamine-
dependent activation of NF-κB signaling in aged perilymphatic
mesenteric tissues [119]. The mast cell/histamine/NF-κB axis
through a myriad of the NF-κB-controlled vasoactive cytokines
induces and maintains aging-associated alterations of the contractil-
ity of neighboring MLVs. However, the underlying molecular mech-
anisms have not yet been discovered but without a doubt are
85Influences of aging
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complex and require detailed investigations of aged lymphatic con-
tractility through careful discovery-driven research.
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CHAPTER
5
CNS lymphatics in health
and disease
J. Winny Yun
a
, J. Steve Alexander
a
,
Felicity N.E. Gavins
a,b
a
Department of Molecular and Cellular Physiology, LSU Health
Sciences Center, Shreveport, LA, United States
b
Brunel University London, Kingston Ln, London, Uxbridge,
United Kingdom
Lymphatics of the central nervous system
The lymphatic vasculature, a secondary transport system that
parallels the blood vasculature, is responsible for clearing interstitial
fluid (ISF), immune cells (lymphocytes, antigen-presenting cells),
and extracellular proteins carried in it, which maintains overall tis-
sue water and solute balance, homeostasis, and immunity [1]. Such
vessels have remained elusive in the context of the brain, and as
such, compartments of the central nervous system (CNS) have been
historically believed to be devoid of a “conventional” lymphatic sys-
tem. This has remained a perplexing conundrum not only because
the brain is exposed to levels of waste products reflecting the brain’s
high metabolic rate, which critically depends on such clearance, but
also because resident cells such as neurons are extremely sensitive to
their ionic changes. Recently, it has been proposed that in the brain,
the clearance of extracellular fluids and extracellular proteins take
on alternative mechanisms that differ from the rest of the organs
of the body. To date, three sets of brain lymphatic clearance path-
ways have been described: (i) the perivascular pathway, (ii) the
93Lymphatic Structure and Function in Health and Disease
https://doi.org/10.1016/B978-0-12-815645-2.00005-8
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