First published online February 1, 2008
Journal of Experimental Biology 211, 510-523 (2008)
Published by The Company of Biologists 2008
doi: 10.1242/jeb.008755
Production of different phenotypes from the same genotype in the same environment by developmental variation
Günter Vogt1,*,
Martin Huber2,
Markus Thiemann3,
Gerald van den Boogaart4,
Oliver J. Schmitz3 and
Christoph D. Schubart2
1 Zoological Institute and Museum, University of Greifswald,
Johann-Sebastian-Bach-Straße 11/12, D-17487 Greifswald, Germany
2 Biology 1, University of Regensburg, Universitätsstraße 31, D-93040
Regensburg, Germany
3 Department of Analytical Chemistry, University of Wuppertal,
Gauss-Straße 20, D-42119 Wuppertal, Germany
4 Department of Mathematics and Computer Science, Jahnstraße 15a,
University of Greifswald, D-17487 Greifswald, Germany
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Fig. 1. Variation of development and growth in juveniles and adolescents. (A,B)
Rather synchronous development of lecithotrophic stage-2 juveniles (arrow)
from dam A on the maternal pleopods (A) and in net culture (B). Arrowhead in B
denotes yolk reserves. Scale bars, 3 mm. (C,D) Size differences among feeding
batch-mates removed from brooding dam A at day 35 after hatching: the juvenile
in C is in stage 3 whilst its sib in D is already in stage 5. Markers in D
denote traits and organs investigated in juveniles: carapace length (black
line), total length (white line), outer branch of 1st antenna (arrowhead)
bearing the olfactory aesthetascs, and chelae of pereiopods 1, 2 and 3
(arrows) bearing the gustatory corrugated setae. Scale bars, 2 mm. (E)
Development of batch-mates (S1–S12) from dam
B5 raised individually in a 12-well microplate from late
embryogenesis to juvenile stage 5 (Juv 5). Development is rather uniform in
embryos and non-feeding stages 1 and 2 but becomes heterogeneous after onset
of feeding in stage 3. (F) Growth differences of five juveniles from dam B
that were size-matched in stage 6 and then cultured in an aquarium without
shelter. 34 days later, one specimen was in stage 11, one in stage 9 and three
were in stage 7. Scale bar, 4 mm. (G) Temporal development of size variation
in eight adolescent batch-mates (B1–B8) from dam B
reared communally under aquarium conditions. Specimens are numbered according
to sequence of egg-laying, which started in B1 at day 158 after
hatching.
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Fig. 2. Variation of growth and reproduction in adults. (A) Temporal development of
variation of carapace length (CL), plotted as coefficient of variation, in
batch-mates from dam B3-1 that were divided into groups of ten
(groups C and D) in the first day of stage 3 (before onset of feeding) and
then kept under identical aquarium conditions. Figures on bars indicate number
of specimens. Arrows indicate time (days) of first egg laying in group. (B)
Variation of mass among batch-mates in groups C and D at day 164 of life. Note
difference in ranges of variation and coefficients of variation (CV) between
groups. (C) Variation in growth and reproduction of communally reared
batch-mates (B1–B7) from dam B. Red vertical bars
indicate time of oviposition. Note position changes of individuals in group
with time. (D) Growth and reproduction of batch-mates
(F1–F4) from dam B3-1 reared together
for 85 days and then kept individually under identical environmental
conditions. Red vertical bars indicate time of oviposition. Arrows denote
appearance of glair glands.
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Fig. 3. Variation of coloration. (A,B) Body areas chosen in juveniles (A) and
adults (B) for comparison of coloration. Frames indicate dorsal posterior (A)
and posterio-lateral part of the carapace (B). (C–F) Areola (marked by
white lines) of four stage-3 batch-mates, showing marked differences of
pigmentation among individuals. (G–J) Pigmentation pattern of same
specimen in juvenile stages 2 (G), 3 (H), 4 (I) and 5 (J), indicating that an
existing pattern is not abruptly changed by moulting but gradually elaborated
by addition of chromatophores. (K–M) Colour pattern of posterio-lateral
carapace area of same adult at time of first reproduction (K), 124 days and 2
moults later (L), and 642 days and 6 moults later (M). The marmoration motifs
are enlarged and modified with time but are still recognizable after more than
21 months. Scale bars, 2 mm. (N–Q) Colour patterns of posterio-lateral
carapace area of dam B (N) and three mature daughters of the same batch
(O–Q). Note striking differences in marmoration among all
individuals.
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Fig. 4. Variation of sense organs. (A) Scanning electron micrograph of olfactory
aesthetascs in an adolescent. (B) Scanning electron micrograph of gustatory
corrugated setae in an adolescent. (C) Means and ranges of the numbers of
aesthetascs per specimen, plotted per stage (Juv 2 to Juv 6) and batch (from
dams A and B and daughters B1 and B3). Juveniles were
removed from the maternal pleopods in stage 2 and then reared under identical
conditions in net culture systems. Shaded areas indicate ranges of all
batches. *Only three individuals analysed in these groups. (D)
Means and ranges of the numbers of corrugated setae per specimen (same
specimens as in C), plotted per juvenile stage and batch. Note differences in
variation among aesthetascs in C and corrugated setae in D. (E) Means and
ranges of the numbers of corrugated setae on pereiopods 1, 2 and 3
(P1–P3), plotted per juvenile stage. Small circles and thin bars
represent offspring of a single female (dam B) and large circles and thick
bars pooled data of all batches (from dams A, B, B1 and
B3).
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Fig. 5. Geometric standard deviations sg=exp{sd[log(x)]} of
corrugated setae (A), aesthetascs (B) and carapace length (C) with 95%
confidence limits. Data are split according to batches (A, B, B1,
B3) and juvenile stages (J2, J3, J4, J5, J6). The normality
approximation in s.d. confidence limits does not hold for those aesthetasc
data sets (in B) that showed no variation, and therefore no confidence limits
are given in these cases. The dots correspond to the estimated standard
deviation. sg is a measure of relative variation and is
approximately 1+CV. Note the typically higher variation in batch
B1.
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Fig. 6. Variation of epigenetic markers. (A) Right–left differences of
marmoration pattern on cephalothorax of mature female. (B) Variation of
fluctuating asymmetry (FA) of corrugated setae (CS) and aesthetascs (Ae) in
168 stage-2 to stage-6 juveniles (stages indicated by numbers on bars) from
dams A, B, B1 and B3. FA is expressed as % deviation
from perfect symmetry (Sym), either to the right (+) or to the left (–).
Red horizontal bars indicate mean FAs of batches. N, number of
individuals investigated per batch. Note independence of fluctuation of both
traits during ontogeny and variation among batches. (C) Alteration of carapace
length (CL), number of aesthetascs and number of corrugated setae in a single
crayfish through nine life stages. Development of the three traits is not
strictly correlated and FA of the sense organs can fluctuate between body
sides. A1-r and A1-l, right and left 1st antennae; P1-r to P3-l, right and
left pereiopods 1, 2 and 3. (D) Variation of body mass and global DNA
methylation of the hepatopancreas and abdominal musculature in three
communally reared adult batch-mates (B2, B7,
B8) at the age of 626 days (left) and four adolescents
(E1–E4) at the age of 188 days (right). Each DNA
sample was measured at least 20 times. Bars indicate +s.d. Variation of mass
and global DNA methylation is apparently not correlated.
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© The Company of Biologists Ltd 2008
