Principles & Practice of Stocking

Overview of potential genetic impacts..

Philip McGinnity

Glasgow

28 November, 2013

Outline What effect does stocking have on wild populations and their
ability to adapt to environmental change?

Theoretical considerations (basic); Identify and map out some of the potential effects (genetic) on wild populations (aid clarity);

Identify useful literature on the subject;

Present results from experiments in the wild that might be helpful (empirical data) in predicting impacts;

Summary;
Context of protecting wild, naturally, self-sustaining salmon populations as a public good in perpetuity.

Theory

Purdom (2001). Trout wars. Trout News, CEFAS, 32, 16-18.

Qualify (a) genetic analysis is a conservative (poor) representation of population

phenotypic (adaptive trait differences) divergence Edmands (2002). TREE 17 (11), 520-527

Easiest way to think of local adaptation is in terms of phenotypic traits, which have a genetic basis (heritable) and provide best match for a given environment (physical; chemical; biological) freshwater, transitional, sea.

Genetic & life history complexity in Atlantic salmon

100 90 80 70
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% Run

60 50 40 30 20

20 10 0 APRIL MAY JUNE JULY AUGUST SEPT

10 0 APRIL MAY JUNE JULY AUGUST SEPT

45 40

100 90
% Run

35 30 25 20 15 10 5 0 APRIL MAY JUNE JULY AUGUST SEPT

80 70

% Run

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Dillane et al. (2008) Molecular Ecology

Theory

Natural selection [process of evolution] in the wild in response to environmental variability [or could be genetic changes underlying phenotypic traits)] imposes demographic costs [reduced abundance] (Burger and Lynch, 1995 Evolution)

Novel environment (TIME)

Gomulkiewicz & Holt 1995 Evolution

Theory
Degree of local adaptation
(match between phenotype + environment)

Once-off stocking

Continual stocking

Perfect adaptation
Increasing maladaptation

Population abundance
(of naturally spawning fish)

Time

Theory

Fisheries Induced Evolution (FIE)

Legacy of the fishery

Think about hatchery effects question in terms of (FIE).

What is the potential for hatchery mediated evolution (HME), i.e. to give rise to genetic change that has (short-term and possibly long-term) fitness consequences, and its impact on the productivity and resilience of individual local populations?

Map - Stocking has potential to affect the both the environment

and the traits expressed..

Evolution (genetic change) in the hatchery environment (Stage 1)

Reduction in genetic variation (sampling error/ founder effect)

Waples & Do (1994). Can. J. Fish. & Aquat. Sci., 51 (Suppl. 1), 310-329

Inbreeding depression

(single event of close inbreeding: 78% reduction in marine survival)

Relaxation of sexual selection

These results indicate that sexual selection favours increased body size and integument coloration in males as well as increases genetic diversity at the MHC by female mate choice

Relaxation of natural selection

Domestication selection

Single generation captive steelhead showed: fitness in hatchery environment fitness in the wild

50%

Domestication selection What traits are affected?

Risk taking; egg size, fecundity, aggressiveness, scheduling of maturation etc.

Reduction in recruitment performance of recipient wild pop 10 Steelhead S/R studies

Chilcote (2003). Can. J. Fish. Aquat. Sci.60: 1057-1067 Chilcote et al. (2011). Can. J. Fish. Aquat. Sci. 68: 511-522 Young (2013), Fisheries Management & Ecology, 20, 434-444.

Long-term monitoring

(Burrishoole River total counts 1969-2009)

Speed of mal-adaptation for life in wild

Araki et al. 2007. Science (also Christie et al. 2013. PNAS 109 (1) 238-242).

Release of hatchery fish into wild (Stage 2)

Replacement (.if a captive brood-stock program is

successful, virtually all of the genes of the post-supplementation population will be derived from fish taken into the hatchery for broodstock Waples & Do, 1994)

Release of hatchery fish into wild

Hybridisation

Hybrid vigour Outbreeding depression (extrinsic)

Outbreeding depression (intrinsic)

Adapted from Burton et al. (2013). Annu. Rev. Ecol. Evol. Syst., 44:14.114.22

Experimental evidence for outbreeding depression

Genetic studies on impact of farm escapes

Empirical evidence - Impact of farm escapes (10 years later)

McGinnity et al. (2003) PRSB

Additive intermediate fitness of hybrids (diagnostic extrinsic Outbreeding response) Heterosis (hybrid vigour not over entire life cycle)

Fitness variation between neighbouring populations

(scale of local adaptation)

50Km
McGinnity et al., 2004 J. Fish Biol.

7 out of 8 traits

(outbreeding depression)

OToole (PhD thesis, unpublished)

Modifying factors on gene flow from hatchery

into wild population

Relative size of introduction; Origin of introduced material (evolved in a different

environments);

Behaviour (sexual selection); Behaviour freshwater experience (life history stage fish
released);

Status of wild population (stable, threatened,

endangered);

Environmental context (change) rate and magnitude;

Carrying capacity (density dependence); Mature male parr (facilitating gene flow);

Mature male parr facilitate gene flow

Hindar et al. (2007) ICES J. Mar. Sci. 63, 1234-1247

w
F

H W

Stocking (broad church)

Modified from Fraser (2008) Evolutionary Applications, 1, 585-586

Genetic Rescue (counter inbreeding depression in wild match historical environments) & Assisted Gene Flow (match future environments)

Houde et al. (2011). Evolutionary applications 4, 634-647

Summary - What effect does stocking have on wild populations?

Domestication selection; reduction of genetic variation via sampling error; inbreeding depression; relaxation of natural selection; relaxation of sexual selection are all potential drivers of evolution of wild populations brought into the hatchery; Compelling evidence that substantial and rapid evolution (genetic change) in the hatchery mal-adapts stocked fish for life in the wild; Marine phase particularly sensitive; Hatchery fish can potentially impact on both the environment (indirect genetic changes) into which they are released & fitness (hybridisation & outbreeding depression) of the recipient population; Introduction of hatchery fish into wild has potential to impact on short-term productivity/yield and possibly long-term (legacy - replacement) of individual population???? Evolution (genetic change) by natural selection can re-establish new equilibrium and offset impacts on yield; recurrent intrusions will continue to depress populations; Hatchery can mediate loss of unique diversity by extinction, replacement, mass hybridisation;

Conclude that HME (genetic change) is an additional stressor on natural populations (overfishing etc); rather than reliever of stress;
Like fishing the magnitude of the stress is relative to the size of the activity and the propensity for gene flow and frequency of introductions.

Summary - What effect does stocking have on ability of wild populations to

adapt to environmental change?

Dont want to lose diversity in period of change

Beaufort Marine Research Award in Fish Population Genetics

Acknowledgement:

This Beaufort Marine Research Award is carried out under the Sea Change Strategy and the Strategy for Science Technology and Innovation (2006-2013), with the support of the Marine Institute, funded under the Marine Research Sub-Programme of the National Development Plan 20072013.