Wb. Campbell et al., THERMALLY-INDUCED CHRONIC DEVELOPMENTAL STRESS IN COHO SALMON - INTEGRATING MEASURES OF MORTALITY, EARLY GROWTH, AND DEVELOPMENTAL INSTABILITY, Oikos, 81(2), 1998, pp. 398-410
Developmental stability, or homeostasis, facilitates the production of
consistent phenotypes by buffering against stress. Fluctuating asymme
try is produced by developmental instability and is manifested as smal
l random departures from bilateral symmetry. Increased fluctuating asy
mmetry is thought to parallel compromised fitness, in part, because st
ress promotes energy dissipation. Compensatory energy expenditures wit
hin the organism are required to complete development, thus promoting
instability through reductions in homeostasis. Increased heterozygosit
y may enhance developmental stability by reducing energy dissipation f
rom stress through increased metabolic efficiency, possibly by providi
ng greater flexibility in metabolic pathways. Traditionally, fluctuati
ng asymmetry has been used as a bioindicator of chronic stress, provid
ed that selective mortality of less fit individuals did not reduce str
ess-mediated increases in fluctuating asymmetry to background levels p
roduced by natural developmental error, or create data inconsistencies
such as higher asymmetry in groups exposed to lower stress. Unfortuna
tely, absence of selective mortality and its effects, while often assu
med, can be difficult to substantiate. We integrated measures of early
growth, mortality, fluctuating asymmetry (mandibular pores, pectoral
finrays, pelvic finrays, and gillrakers on the upper and lower arms of
the first branchial arch) and directional asymmetry (branchiostegal r
ays) to assess chronic thermal stress (fluctuating temperatures as opp
osed to ambient temperatures) in developing eggs from two different co
ho salmon (Oncorhynchus kisutch) stocks and their reciprocal hybrids.
Hybridization provided insight on the capacity of heterozygosity to re
duce stress during development. Although egg losses were consistently
higher in crosses exposed to fluctuating temperatures, egg mortality w
as predominantly a function of maternal stock of origin. Post-hatch lo
sses were higher in crosses exposed to ambient temperatures than in cr
osses exposed to fluctuating temperatures during embryogenesis. Observ
ed patterns of early growth revealed no heterosis. but instead reflect
ed maternal effects, with some crosses slowing growth and yolk utiliza
tion when exposed to fluctuating temperatures. Analyses of fluctuating
asymmetry also showed no effects from heterosis. While analyses of co
mposite asymmetry scores and branchiostegal rays were inconclusive, an
alyses of individual characters showed significantly higher fluctuatin
g asymmetry in pelvic finray counts and a marginal change in the numbe
rs of fish asymmetric for this character in crosses exposed to chronic
thermal stress. In contrast, the fluctuating asymmetry. in lower gill
raker counts was significantly higher in crosses exposed to ambient te
mperatures and there were significantly more fish asymmetric for this
character. Data on mortalities and fluctuating asymmetry indicate pelv
ic finray development was thermally stressed, while the heightened flu
ctuating asymmetry in lower gillraker counts under ambient temperature
s was due to a greater frequency of less fit fish that had not been cu
lled by thermal stress. Changes in early growth patterns in response t
o developmental stress yielded no parallel responses in meristic chara
cters. We conclude that chronic thermal stress produced both selective
ly lethal and sublethal effects that directly shaped fluctuating asymm
etry and fitness profiles in these crosses. Implicit in this conclusio
n is that developmental instability analyses can detect more than just
chronic sublethal stress, thus providing substantial credence for usi
ng instability studies as proactive bioassessment methodologies.