Understanding the biological underpinnings of fragile
X syndrome
Pietro Chiurazzi
a
, Giovanni Neri
a
and Ben A. Oostra
b
Purpose of review
The purpose of this review is to present the latest findings on
fragile X syndrome and to put them into perspective. Fragile X
syndrome is a relatively common form of inherited mental
retardation, caused by loss of function of the FMR1 gene on
the long arm of the X chromosome. The molecular
mechanisms underlying the syndrome are complex and
continue to surprise researchers more than 12 years after the
cloning of the gene.
Recent findings
We will specifically discuss the various aspects of the clinical
phenotype, reassessed with the employment of functional
imaging and electrophysiological techniques. The unexpected
finding of a pathologic phenotype in premutation carriers is
highlighted, as it represents a new and distinct condition with
a different presentation in males and females. The third section
deals briefly with the various functions of the FMRP protein, an
RNA-binding protein interacting with multiple RNA molecules
as well as proteins. It is important to realize that FMRP is
probably changing partners several times, depending on its
localization, on posttranslational modifications and on the
available interacting proteins. In the following section, we
present in short recent discoveries on the defective neuronal
circuits in the fragile X syndrome. Most of these new data were
made available by the study of animal models, mostly the Fmr1
knockout mouse, but also Drosophila.
Summary
We briefly discuss the alternative options for treating fragile X
syndrome. Presently, a neuropharmacological approach acting
on either critical receptors or aimed at reactivating the silenced
FMR1 gene appears promising.
Keywords
fragile X syndrome, mental retardation, translational repression,
synaptogenesis, neuronal plasticity
Curr Opin Pediatr 15:559–566. © 2003 Lippincott Williams & Wilkins.
Introduction
In the past 2 years, several advances have been made in
the understanding of fragile X syndrome, a heritable
form of mental retardation caused by mutation of the
X-linked FMR1 gene [1]. The characteristics of FMRP,
the protein encoded by FMR1, and of its related FXR
proteins are gradually being revealed [2,3]. FMRP inter-
acts with multiple RNA molecules and proteins at the
same time [4,5], while shuttling between the nucleus
and the cytoplasm. In neurons, the cells in which FMR1
is expressed at the highest level, FMRP reaches post-
synaptic terminations in the dendrites by active trans-
portation as part of a ribonucleoprotein complex
(mRNP), bound to a kinesin motor and following cyto-
skeletal microtubules and actin filaments [5]. At the syn-
apse, FMRP acts as a negative regulator of translation
[6,7], possibly switching between active and inactive
statesdependingonitsphosphorylation[8•],onitsbind-
ing partners, and on the activation of specific receptors
[9].TheabsenceofFMRPinfragileXpatientsaswellas
in animal models causes both functional and morpho-
logic changes of synapses [9,10]. In Drosophila, an indi-
rect connection has been found between FMRP and
Rac1 GTPase via the interacting protein CYFIP [11],
linking FMRP to a molecular pathway crucial for cyto-
skeleton remodeling [12].
However, the most surprising recent finding has been
the discovery of a distinct phenotype in male carriers of
a premutation: these intellectually normal individuals
have increased FMR1 mRNA levels and may develop
tremor, ataxia, and dementia with aging [13], usually ac-
companied by extensive brain atrophy. In fact, this neu-
rodegenerative condition, the fragile X premutation
tremor/ataxia syndrome (FXTAS), is entirely distinct
from fragile X syndrome, and its cause appears to be the
accumulation of premutated FMR1 mRNA molecules
rather than their absence.
Clinical aspects, behavior, and diagnostics
Fragile X syndrome is clinically characterized by a long
and relatively narrow face, large protruding ears, mac-
roorchidism, and joint hyperlaxity. A recent report has
statistically compared physical measurements of patients
and unaffected members of their families in a large set
fromAustraliaandtheUnitedStates[14],confirmingthe
significance of the above-mentioned anomalies and
documenting also a decreased body height and limb
length. The authors of this report suggest that a possible
a
Institute of Medical Genetics, Catholic University, Rome, Italy; and
b
Department of
Clinical Genetics, Erasmus University, Rotterdam, The Netherlands
Correspondence to Ben A. Oostra, PhD, Department of Clinical Genetics, Erasmus
University, PO Box 1738, Rotterdam, 3000 DR, The Netherlands
E-mail: b.oostra@erasmusmc.nl
Current Opinion in Pediatrics 2003, 15:559–566
Abbreviations
FXTAS fragile X premutation tremor/ataxia syndrome
KO knockout
POF premature ovarian failure
© 2003 Lippincott Williams & Wilkins
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