1 3
J Biol Inorg Chem
DOI 10.1007/s00775-015-1281-3
ORIGINAL PAPER
Elemental mapping of the entire intact Drosophila gastrointestinal
tract
Michael W. M. Jones
1,2
· Martin D. de Jonge
1
· Simon A. James
1
· Richard Burke
3
Received: 21 May 2015 / Accepted: 23 June 2015
© SBIC 2015
Malpighian tubules. This work simultaneously reveals dis-
tribution of a number of biologically important elements
in entire, intact GI tracts. These distributions revealed not
only a previously undescribed Ca/Mn co-localization, but
also the unexpected presence of additional Cu accumula-
tions in the iron cell region.
Keywords Drosophila · Metal ion absorption ·
Gastrointestinal tract · Synchrotron · X-ray fluorescence
microscopy
Introduction
Essential biometals such as copper (Cu), iron (Fe) and zinc
(Zn) play numerous roles in animal growth, development
and maintenance, acting as structural or catalytic co-factors
for collectively hundreds of proteins. The absolute require-
ment for these dietary nutrients is reflected in the embry-
onic lethality caused by mutation of mouse ion transporter
genes such as Ctr1 (Cu uptake [1, 2]) and Zip4 (Zn uptake
[3]) and the severe iron deficiency anaemia that results
from deletion of divalent metal transporter 1 (DMT1, [4]).
Biometals are selectively absorbed from the dietary con-
tents of the gastrointestinal (GI) tract into the bloodstream
by active transport across the apical (lumenal) and basolat-
eral (circulatory) membranes of the intestinal enterocytes.
For the most part, the metal transport proteins required for
this absorption are well characterized. For example, Ctr1
is needed for uptake of Cu into the enterocytes [5] and
ATP7A mediates Cu efflux across their basolateral mem-
brane into the bloodstream [6], while for Zn, these func-
tions are served by the Zip4 [7] and ZnT1 [8] proteins,
respectively. Additionally, three pathways act to import
dietary Fe into the enterocytes: uptake of heme by an as yet
Abstract The main role of the animal gastrointestinal
(GI) tract is the selective absorption of dietary nutrients
from ingested food sources. One class of vital micronutri-
ents are the essential biometals such as copper, zinc and
iron, which participate in a plethora of biological process,
acting as enzymatic or structural co-factors for numerous
proteins and also as important cellular signalling mole-
cules. To help elucidate the mechanisms by which biomet-
als are absorbed from the diet, we mapped elemental dis-
tribution in entire, intact Drosophila larval GI tracts using
synchrotron X-ray fluorescence microscopy. Our results
revealed distinct regions of the GI tract enriched for spe-
cific metals. Copper was found to be concentrated in the
copper cell region but also in the region directly anterior to
the copper cells and unexpectedly, in the middle midgut/
iron cell region as well. Iron was observed exclusively in
the iron cell region, confirming previous work with iron-
specific histological stains. Zinc was observed throughout
the GI tract with an increased accumulation in the pos-
terior midgut region, while manganese was seen to co-
localize with calcium specifically in clusters in the distal
Electronic supplementary material The online version of this
article (doi:10.1007/s00775-015-1281-3) contains supplementary
material, which is available to authorized users.
* Richard Burke
richard.burke@monash.edu
1
Australian Synchrotron, 800 Blackburn Road, Clayton 3168,
Australia
2
ARC Centre of Excellence for Advanced Molecular Imaging,
Australian Synchrotron, 800 Blackburn Road, Clayton 3168,
Australia
3
School of Biological Sciences, Monash University,
Wellington Road, Clayton 3163, Australia