Growth and Reproduction in the Deep Sea

1
Growth and Reproduction in the Deep Sea

• Rachel Whitney
• Brad Morris

2
Ecological constraints of Growth and Reproduction

• Environmental constraints must be evaluated,
  since temperature and salinity in the deep sea
  rarely fluctuate, and since light is absent in
  benthic zones, then food must be the predominant
  constraint on growth and reproduction based on an
  increased availability of nutrients in the
  sediment layers
• 2 major ecological factors
• Benthic storms and turbidity currents
• Particle flux
• Others related to habitat
• Hydrothermal vent dynamics
• Ephemeral and temporary habitats

3
Benthic storms and turbidity flow through
microenvironments

• Horizontal advection with seasonal variance
• Seasonal changes in the benthic boundary layer
  involved in the production of high laminar flow
  rates in the nephiloid layer
• Rates as low as 7 cm s-1 (just 1 m off bottom)
  can disturb benthic settlement and promotes
  resuspension events in the benthos
• Epifaunal and infaunal organisms can disturb
  settlement and promote resuspension through
  activity

4
Particle Flux

• Seasonal variation
• Rate of particulate sink can vary greatly from
  1-1000 m per day
• About 90 of sinking particulate matter is
  consumed before reaching the benthos, with
  roughly 1 eventually settling on the bottom
• Periodicity of particle flux is easier to
  determine above the BBL.
• Diurnal tide flux can cause elevation of
  particulate matter in the water column

5
Seasonal signals

• Small particulate organic matter entering the
  BBL is the main source of food for many
  organisms living there JD Gage
• Varies with proximity to landmasses and
  freshwater offshoots
• Seasonal signal can be significantly diminished
  in oligotrophic seas, however deep trenches and
  canyons off coastal waters can show an enhanced
  localization of particulate flux due to shelf
  isolation

6
Food Falls

• Large animal
• Macroalgae and seagrasses
• Phytodetritus

7
Growth

• Measuring techniques
• Strategies
• Controls

8
Measuring Growth

• Growth bands in calcareous plates of echinoderms
• Coarse-pored areas reflect periods of relatively
  rapid growth, fine-pored areas created when
  growth is slower
• Growth rings in otoliths of fish
• Direct observations

9
Wilson (1988)
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Growth Strategies

• Decreasing growth rate with increasing age
• Modeled by Von Bertalanffy
• LtL8-( L8-L0)e-K(t-t0)
• Lt size at time t
• L8 max size attainable
• L0settlement size
• T0settlement time
• Krate constant

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Growth Strategies

• Growth rate increases initially, and then
  decreases with age
• Modeled with Gompertz function
• LtL8e-e-K(t-t0)
• Lt size at time t
• L8 max size attainable
• t0settlement time
• Krate constant
• Common in urchins

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• Gage, J.D. (1990)

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Growth Strategies

• Indeterminate growth growth and reproduction
  occur simultaneously.
• Ex. Echinus affinis After maturity the volume
  rate of growth does not decline but becomes
  constant.

14
Middleton and Gurner (1998)
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Growth Strategies

• Sexual dimorphism
• Ex. Red crab, Chaceon affinis
• Males grow larger than females
• Secondary sexual characteristics related to
  reproductive behaviours

Fernández-Vergaz et al (2000)
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Growth Strategies

• Growth Phases
• Crustaceans have 2 growth phases puberal and
  mature.
• In the crab, morphometric maturity is independent
  of physiological maturity.
• Chaceon affinis females become morphometrically
  mature before they become physiologically mature,
  but males attain sexually mature before they
  reach morphometric maturity

17
Fernández-Vergaz et al (2000)
18
Controls of Growth

• Depth
• Growth rate of two upper bathyal Echinus sp were
  between those of the faster-growing shallow-water
  Echinus esculentus, and the slower-growing
  deep-sea Echinus affinis. Gage et al 1986)
• Lampitt (1990) calculated growth rates in
  deep-sea barnacles several times lower than rates
  reported for near-surface barnacles.
• Sumida et al (2000) found that brittle star
  Ophiocten gracilis post larvae collected outside
  their depth range (deeper) showed slower rates of
  growth.

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Controls of Growth

• Food availability
• Some small deep-sea species can accelerate growth
  and experience early maturation when food
  availability is increased.
• Ex. Barnacle, Poecilasma kaempferi studied by
  Lampitt (1990) showed increase in growth rate by
  several times during mid-May phytodetritus
  deposition from spring bloom.
• Varying feeding rate in lab can induce growth
  zones in calcareous test of urchins.

20
Lampitt (1990)
21
Controls of Growth

• Seasonal patterns
• Patterns in growth bands in skeletal plates of
  deep-sea echinoids have been thought to reflect
  seasonal variability in growth rates, maybe a
  result of seasonal cycles in reproduction or
  nutrition.

22
Controls of Growth

• Activity Rhythms
• Photoperiod is the zeitgeber that initiates
  activity in shallow water fish. (Daily growth
  rings seen in otoliths)
• Macrourid sagittae have daily growth rings
  similar to shallow water species
• Wilson (1988) suggested that tidal cycles could
  be the cue in deep-sea fish, since they can sense
  the tidal cues and current directions.

23
Reproduction

• Form
• Pattern
• Dependance
• Adaptation

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Forms

• Two allocations
• Gonochoristic
• Hermaphroditic (teratogenic in some species)
• A. carchara gonochoric ophuroid, over 40 spp.
  of Ophuroids are hermaphroditic
• Species specialization

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Patterns of reproduction

• Slow egg producing
• Seasonal (periodic continuous)
• Continuous (continuous asynchronous)
• Auto or heterosynthetic vitellogenesis
• Fast egg producing
• Oppourtunistic
• Modified heterosynthetic vitellogenesis

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Reproductive Patterns
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Vitellogenesis and Phylogenic constraints

• Yolk production in the female of a gonochoristic
  spp.
• Does not fit any one pattern of reproduction, no
  discrete form, however varies across taxa and
  more similar in closely relate spp.
• Fecundity increase with increase in nutrients,
  phylogenetically dependant
• Critical Nutrient Hypothesis
• Annual and monthly breeders different

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Semelparity vs. Iteroparity

• Volitile vs. Stable habitat
• Seasonal vs. Continuous breeder
• - ephemeral habitats show environmental
  fluctuation and therefore background species
  frequently displaced
• - oligotrophic areas generally indicative of
  more established assemblages

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Adaptations for Reproduction

• Given by
• Temporal patterns of Reproduction
• Fecundity and Fertility
• Breeding and Development (reviewed later in this
  class)

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Periodicity in Gametogenesis

• Quasi-continuous/periodic
• Synchrony between individuals/ under
  environmental or endogenous control
• Gametogenesis
• Instantaneous fecundity
• Maximum oocyte size
• Seasonality of development

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Possible Mechanisms of Seasonal Reproduction
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Examples from comparative species

• Protobranch bivalves
• M. cuneata (continuous), L. pustulosa, Y.
  jefferysi (seasonal)
• Asteroids
• H inermis, S. Chuni and S horridus (all
  continuous)
• Ophuroids
• O. ljungmani

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Fecundity and Fertility

• Different for semelparous and iteroparous species
• Depends on type of development of species as well
  which in turn indicates the overall number of
  oocytes per gonad
• Mature individuals in reproductive state will
  allocate more energy expenditure to gametogenesis
  at somatic expense

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Examples in fecundity

• Ophuroids
• A.carchara embyros in broding females from 1 to
  72, low fecundity
• O. Graciallis disk dependant fecundity (up to
  ca. 51000)
• Protobraches
• M.cuneata lowest fecundity with largest oocyte
  size
• L.pustulosa and Y.jefferysi higher fecundity with
  significantly decreased egg size

35
Restrictions to evaluation of Growth and
Reproduction

• Growth and reproductive measures lacking for many
  species
• Collection methods skewed
• Indirect calculations, hypothesis testing
• Comparative coastal to deep sea lifestyles

36
References

• Eckelbarger, K.J., Watling, L., 1995. Role of
  phylogenetic constraints in determining
  reproductive patterns in deep-sea invertebrates.
  Invertebrate biology 114 (3) 256-269.
•  
• Fernandez-Vergaz,-V. Lopez-Abellan,-L.J.
  Balguerias,-E., 2000.Morphometric, functional and
  sexual maturity of the deep-sea red crab Chaceon
  affinis inhabiting Canary Island waters
  Chronology of maturation. Mar-Ecol-Prog-Ser 204
  169-178
• Gage, J.D. 1990.  Skeletal growth markers in the
  deep-sea brittle stars Ophiura ljungmani and
  Ophiomusium lymani. Marine Biology 104, pp
  427-435
• Gage, J.D., 1994. Recruitment Ecology and Age
  structure of Deep -Sea Invertebrate populations.
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• Gage, J. D., Tyler, P.A., and Nichols, D.
  Reproduction and growth of Echinus acutus var
  norvegicus Duben  Koren and E. elegans Duben
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•  
• Harrison, K. 1988. Seasonal reproduction in
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•  
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•  
• Lampitt,-R.S., 1990. Directly measured rapid
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• Lightfoot, R.H., Tyler, P.A., and Gage, J.D.
  1979. Seasonal reproduction in deep-sea bivalves
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•  
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•  
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•  
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• Tyler, P. A., Campos-Creasey, L.S., and Giles, L.
  A., 1994. Environmental control of
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•  
•  

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