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The Journals of Gerontology Series A: Biological Sciences and Medical Sciences 62:1361-1363 (2007)
© 2007 The Gerontological Society of America


COMMENTARY ON A RECENT PUBLICATION

Age Structure and Senescence in Long-Term Cohorts of Eisenia andrei (Oligochaeta: Lumbricidae)

Christian Mulder, Jan Hendriks, Rob Baerselman and Leo Posthuma

1 National Institute for Public Health and the Environment, Bilthoven, The Netherlands. 2 Department of Environmental Science, Radboud University, Nijmegen, The Netherlands.

Address correspondence to Dr. Christian Mulder, Laboratory for Ecological Risk Assessment, National Institute for Public Health and the Environment, P. O. Box 1, NL-3720BA Bilthoven, The Netherlands. E-mail: christian.mulder{at}rivm.nl

CHEN and colleagues (1) discussed fitness and aging relationships, considering senescence evolution in laboratory lines of nematodes. They concluded that fitness costs for Caenorhabditis elegans are consistent with antagonistic pleiotropy. Fitness and demography might be dominated by indirect genetic effects in other organisms too. We provide an additional explanation for senescence evolution by allometric mass-specific costs, as shown by long-term dynamics of reproduction, growth, and mortality of Eisenia andrei age cohorts. Empirical results collected during a 9-year experiment are provided and discussed. In this context we introduced mass-derived Gompertzian Metabolic Rates (GMR), which are benchmarks in comparative physiology.

Changes in the culture conditions affect the progeny of invertebrates, as shown by selecting populations under modeled situations in the laboratory (2), but in contrast to nematodes (3,4), concrete estimates of longevity for most invertebrates are lacking. Recently, Chen and colleagues (1) rejuvenated the discussion on fitness estimates of worm-like organisms. Our long-term study on the senescence (increase of mortality rate with age) of the earthworm Eisenia andrei supports other conclusions on mortality rates of invertebrates born and kept in captivity. The median longevity of E. andrei in the artificial OECD (Organisation for Economic Co-operation and Development, www.oecd.org) soil is 63 months, the median survivorship 240 weeks. The entire life span at 18°C of the oldest specimen was 8.73 years. E. andrei showed an individual average cocoon production of 0.8–2.0 cocoons per week. As in a study on the closely related Eisenia foetida (5), these cocoons were pale with a mucousy, jelly-like consistency at birth, although the cocoons of E. andrei took on a much more brownish hue within some hours. For large age cohorts, the mortality rate in laboratory lines from big cocoons reaches a maximum of 29.8% after 42.5 months, and to this age survived 65.0% of the Eisenia adults who survived their birth (Table 1 and Figure 1). Shortly after this event, the decaying oscillations in body mass highly increased ({chi}2 = 101, p <.0001). Therefore, such an unexpected mortality peak can be seen as a bifurcation in our data, where the earthworm population apparently failed to meet some conditions required for their growth. Moreover, the period between weeks 170 and 173 might reflect the threshold between young and old ages previously described (9). Under standard conditions like those of our experiment, body-mass values of animals ranging in size from protists to mammals reflect their metabolism, fecundity, and life span (7). As all data were normalized to constant laboratory temperature, a metabolic rate as function of body mass depicts the maintaining energy costs for keeping the Eisenia body active, thus fit. [Our mass-specific definition of fitness differs in this aspect from that used by Chen and colleagues (1), as they have chosen an integrative stage-specific definition.] These allometric implications clearly show that oscillations in the body-mass values are primary determinants for entire populations. Such a finding is reasonable as this increasing lack of fit strictly reflects the late weeks of Eisenia's life, during which the aging process reaches a finite plateau. Only long-term experimental designs allow evaluation of possible explanations which do not involve a causative role between body and productivity.


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Table 1. Senescence of the Eisenia Age Cohort.

 

Figure 01
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Figure 1. Top: Body-mass M as function of time T (weeks) of the 37 Eisenia andrei adults from small cocoons (left) and the 40 adults from big cocoons (right). Bottom: Gompertzian Metabolic Rate (mL O2 day–1) and the total number of cocoons. In this new biodemographic study, we used the fresh body-mass values of individually maintained E. andrei as predictors of metabolic rate (7), hence fitness and life-span variation (8). Mass-derived GMR fluctuates broadly in synchrony with the cocoons (Pearson's coefficient = 0.776, p <.0001). Arrow: death of 11 earthworms between the 170th and the 173th week. Most oscillations occur after the 170th week (analysis of variance F1,127 = 31.57, p <.0001), but in contrast to other studies (1,4), senescent decline is not consistent with the postreproductive period of life of Eisenia

 
We conclude that the aging process of the investigated earthworms provides suggestions for successive time derivates in more general cases, suggestions that are different from those derived from C. elegans stages (3,4) and comparable to those derived from mammals (6,7).

SUPPLEMENTARY DATASET: EISENIA ANDREI

The Supplementary Dataset is linked to the online article in the December issue at http://biomed.gerontologyjournals.org/content/vol62/issue12/.

Acknowledgments

The authors wish it to be known that, in their opinion, the last two authors should be regarded as joint last authors.

References

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  2. Parsons PA. Evolutionary rates: stress and species boundaries. Annu Rev Ecol Syst. 1991;22:1-18.
  3. Van Voorhies WA, Fuchs J, Thomas S. The longevity of Caenorhabditis elegans in soil. Biol Lett. 2005;1:247-249.[Abstract/Free Full Text]
  4. Johnson TE, Wu D, Tedesco P, Dames S, Vaupel JW. Age-specific demographic profiles of longevity mutants in Caenorhabditis elegans show segmental effects. J Gerontol Biol Sci. 2001;56A:B331-B339.[Abstract/Free Full Text]
  5. Storey KG. Cell lineage and pattern formation in the earthworm embryo. Development. 1989;107:519-531.[Abstract]
  6. Gompertz B. On the nature of the function expressive of the law of human mortality and on a new mode of determining life contingencies. Phil Trans R Soc London. 1825;115:513-585.
  7. White CR, Seymour RS. Does basal metabolic rate contain a useful signal? Mammalian BMR allometry and correlations with a selection of physiological, ecological, and life-history variables. Physiol Biochem Zool. 2004;77:929-941.[Medline]
  8. Watanabe N, Nagamatsu Y, Gengyo-Ando K, Mitani S, Ohshima Y. Control of body size by SMA-5, a homolog of MAP kinase BMK1/ERK5, in C. elegans. Development. 2005;132:3175-3184.[Abstract/Free Full Text]
  9. Li N. Estimating national disability risk. Theor Popul Biol. 2004;65:389-400.[Medline]




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