THE PERCEPTION OF PLANT INVASION  

        The biology of the invasion process as presented in the Invasion Overview section is rational and experimentally tractable.  What is less apparent is the human component of the selection process that creates opportunity spaces into which invasive species disperse.  The direct effects of human activity are also more discernable than the indirect effects.  Of critical importance is the role human perception plays in selection and creation of opportunity space for invasive species.  
   
     A species succeeding in occupying a locality must be perceived by humans as being problematic for it to be labeled invasive.  The perception of a plant species as invasive by humans is a complex, often highly subjective process.  Despite this, there are several systematic ways to understand how human perception and cultural values create selection pressure and opportunity spaces conducive to plant invasions.  They include insights gained from public policy and reflection on human values.  These social and perceptual factors are inherently anthropological and anthrocentric in nature, and need to be understood in those contexts for a complete understanding of the forces of selection conducive to invasion.

anthropology              
1:  the scientific study of human beings, their origins, distribution, physical attributes and culture (Anonymous, 2001)
2:  the study of man, his origins, physical characteristics, institutions, religious beliefs, social relationships, etc. (Anonymous, 1979)

culture             
1:  the skills, arts, etc. of a given people in a given period (Anonymous,
2001)
2:  the entire range of customs, beliefs, social forms, and material traits of a religious, social, or racial group (Anonymous, 2001)
3:  the total of the inherited ideas, beliefs, values, and knowledge, which constitute the shared bases of social action
4:  the total range of activities and ideas of a group of people with shared traditions, which are transmitted and reinforced by members of the group (Anonymous, 1979)

antrocentric               
1:  centering in man (Anonymous, 2001)

anthropocentric          
1:  regarding man as the most important and central factor in the universe (Anonymous, 1979)

Public Policy
   
Public policy can provide a starting point to determine human perceptions of invasive species, an expression of human values.  Of particular interest is public policy on invasive species promulgated by the U.S. Federal government in Executive Order 13112 of February 3, 1999 (Anonymous, 1999, 2004b).  Research, management and dissemination of information about invasive species in the U.S. are funded by government agencies in compliance with this order.  The terminology used in this public policy statement reveals how some perceive invasion biology.  Therein (Anonymous, 1999, 2004b) they define several terms, below included with definitions from more conventional sources:

invasive species         
1:  an alien species whose introduction does or is likely to cause economic or environmental harm or harm to human health (Anonymous, 1999)
2:  a species that is non-native (or alien) to the ecosystem under consideration and whose introduction causes or is likely to cause economic or environmental harm or harm human health (Anonymous, 2004)

alien species               
1:  with respect to a particular ecosystem, any species, including seeds, eggs, spores, or other biological material capable or propagating that species, that is not native to that ecosystem (Anonymous, 1999)
2:  non-native; a species occurring in an area to which it is not native (Lincoln et al., 1998)

native species            
1:  with respect to a particular ecosystem, a species that, other than as a result of an introduction, historically occurred or currently occurs in an ecosystem (Anonymous, 1999)

native                          
1:  relating to the indigenous inhabitants of a country or area; a local
inhabitant; an indigenous plant or animal (Anonymous, 2001)
2:  relating or belonging to a person or thing by virtue of conditions existing at the time of birth; born in particular place (Anonymous, 1979)
3:  indigenous; living naturally within a given area; used of a plant species that occurs at least partly in natural habitats and is consistently associated with certain other species in these habitats (Lincoln et al., 1998)

nativism                      
1:  the doctrine of innate ideas
2:  in U.S., the advocacy of the claim of native as opposed to that of naturalized Americans (Anonymous, 2001)
3:  Chiefly U.S. the policy of favouring the natives of a country over the immigrants (Anonymous, 1979)

natural                        
1:  of or produced by nature (Anonymous, 2001)
2:  in accordance with human nature (Anonymous, 1979)          
3:  not affected by man or civilization; uncultivated; wild (Anonymous, 1979)

introduction                
1:  intentional or unintentional escape, release, dissemination, or placement or a species into an ecosystem as a result of human activity (Anonymous, 1999)  

    Several aspects of invasion biology are revealed in these definitions.  These include the concept of economic, environmental and human harm; the differentiation between alien and native species; the existence of natural conditions; and the purposeful introduction of a plant species to a locality. 
   
The purpose of this paper is only to highlight the explicit statements of human goals and values that may influence invasion biology.  Of specific importance to public policy is the value placed on nativism, natural conditions and the different categories of harm.  How public policy is implemented with these guiding, often subjective, concepts is at the heart of how these species are managed.  The management elicited by public policy is the selection pressure these invasive species will respond and adapt to in their subsequent evolution.

Human Values. 
   
The historical expansion of human populations, and their activities, has affected almost every habitat on earth to some extent, either directly or indirectly.  Air and water pollution alone have affected much of the surface biology of earth (e.g. CO2, O3).  Human perception of what is natural and indigenous, what is disturbed and artificial, is therefore compromised to some degree.  In one form or another, willingly or not, the earth is the garden of humanity.  The equivocal nature of what harm is caused by invasive species is therefore confounded by the heterogeneous array of human viewpoints and aesthetic values of what is desirable in landscapes.  This heterogeneity of opinion is not resolvable but remains at the core of invasion biology because values guide activity and management.  For better or worse, the actualization of human values creates opportunity space for new species to invade:  they are a direct reflection of human activity.
   
The best expression of human-mediated invasion biology can be found in agriculture.  With the advent of agriculture some 10,000 years ago, hunter-gatherer and nomadic peoples were displaced gradually by spatially sedentary agriculturists.  The opportunity space for agriculture was vast.  Humans imposed disturbance regimes on those spaces (e.g. soil tillage) and favored plant species with desirable phenotypic traits to cultivate and harvest.  Evolutionary changes in those cultivated species led to somewhat ironic consequences: the formation of stable, long-lived wild-crop-weed complexes (de Wet, 1966; de Wet and Harlan, 1975).  Wild progenitor species were domesticated.  Crop phenotypes escaped cultivation and developed weedy habits ideal for infestation with their crop relative, and both shared space with the original wild relatives.  Gene flow was continuous between these closely related forms of the same species-group, an ideal genetic situation for the longevity of the species.  Archetypical examples of these wild-crop-weed complexes are found in Amarathus (grain amaranth, pigweeds), Setaria (foxtail millet, green foxtail; Dekker, 2003; 2004b), Brassica (rapeseed and wild mustards), Helianthus (sunflowers), Avena (oat), Oryza (rice), sorghum (crop, johnsongrass), Solanum (potatoes, nightshades), and Hordeum (barley, foxtail barley).  
   
The most important current agricultural plant invasion is the introduction of transgenic crops, often on a vast scale (e.g. glyphosate-resistant crops).  Introduction of any trans-gene into the crop cultivars of these wild-crop-weed complexes increases the chances of introgression into its related non-cultivated weedy and wild phenotypes (e.g. Dekker and Comstock, 1992).  The development of these biotechnologies in wild-crop-weed complexes fulfill the conjecture provided in the introduction: a critical interaction of disturbance, dispersal and plant traits adapted for the resultant opportunity space.  The introduction of such biotechnologies as herbicide-resistant crops provides a mixture of environmental and economic benefit and harm which makes implementation of public policy as defined by U.S. Federal policy (Anonymous, 1999, 2004a) somewhat problematic and highlights the complex interaction of biology and human values.

CONCLUSIONS

"With the present tremendous population explosion the most common habitat has become man-made, and it may not be many centuries before this will be the only habitat available.  With the disappearance of stable habitats, truly wild species will be the first to become extinct.  Wild colonizers may survive as long as habitats remain that are only sporadically disturbed by man.  Eventually these must also disappear and Homo sapiens, the ultimate of all weeds, will lord it over the domain he has created for himself, his companion weeds, his crops and domesticated animals."

-J.M.J. de Wet, 1966

The Human Role in Creating Opportunity Space for Plant Invasion
   
Invasion biology is a reflection of the impact human populations have on the earth's ecology.  Public policy is currently focused on management and control of specific species, but at the same time ignoring the fundamental and complex sources of these changes in biological communities.  Fundamentally the problem is human: human population size and collateral disturbance, human dispersal of invasive species propagules, heterogeneous human values about the nature of harm and beauty, and the priorities of human scientific endeavors.  In all this there may be some benefit to humans by exploiting the very traits we despise most for plant improvement.
Human population size.  Human population size may be the primary cause of invasion biology and changing community structure.  With expanding populations are consequential changes in land use and spatial organization, increasing direct and indirect disturbance, increased resource use and loss, and other changes to habitats.
Global propagule dispersal.  Human global traffic has increased significantly in recent history.  There exists an increased "transferability" of everything in human global society: trade goods and services, human travel and transportation, and ideas.  Swept along with this traffic have been vastly increased opportunities of dispersal of biological propagules into available opportunity spaces.  World grain traffic alone has moved immense quantities of plant propagules over historical times, despite our best efforts to control the more noxious forms.
Human disturbance.  Landscapes and habitats around the world have been influenced by this byproduct of human activity.  Air and water pollution is ubiquitous and affects almost all spaces on the earth's surface.  Direct and indirect disturbances by humans have altered most of these spaces and the ecological relationships in biological communities, leaving vast new opportunity spaces open and available to species with traits allowing their exploitation.
Human values and culture.  Human perception of these changes, public policy initiatives defining environmental and economic harm, and human aesthetic values provide  heterogenous and often conflicting value systems to be compromised in reaching a consensus on the best solutions.  Contributing to the situation is a recent increase in perceived fear of alien invasion, and a nativistic reaction to these fears.
Science of invasion biology.  There is not a meaningful difference between the invasion process and the processes of ecological community assembly, succession and the evolution of niche differentiation by speciation.  Despite this, disciplinary barriers are apparent in the differentiation of invasion biology science in unmanaged and managed habitats: agricultural weed biology and invasive plant biology are often separated in the scientific academy.  Both these realms are unified by disturbance as a prime motivator of change in community structure.  The scale of habitats in time and space is continuous; and all communities are inter-related and inter-dependent.  Agriculturalists often do not completely embrace the invasion process in understanding population shifts, and ecologists studying unmanaged systems often fail to recognize the role of indirect disturbance and dependence on adjacent agricultural habitats in the larger landscape. The science of both will advance when the unifying principles underlying both types of undesirable species are acknowledged in a larger view of invasion biology.
Utilizing and Exploiting Beneficial Plant Invasive Species

weed    1:  a plant whose virtues have not been yet discovered (attributed to R.W. Emerson, 1878).

    Imagine a world in which global warming results in Antarctic ice melting and exposing a terrestrial habitat.  What would the first higher plant invaders look like?  Imagine that humans colonize Mars.  What will their controlled environment plant communities look like?  In either case, does there now exist plant traits that may provide answers to these two speculative questions?  Would exploitation of the traits that allow invasives to succeed provide us with better plant communities, whether Antarctic, Martian, agricultural or scenic? 
   
As humans alter the earth's habitats directly and indirectly, some consideration should be given to preserving and exploiting the germplasm of our best-adapted invasive and weedy species for such a future.  Preservation of weedy and invasive genotypes is an emerging issue in science as novel and unique weedy biotypes are lost with the significant changes biotechnological crop introductions are causing (e.g. Anonymous, 2004a; Marshall et al., 2003).  The traits these noxious plants possess may provide novel solutions to the problems we create in expanding human communities.  The only obstacle to utilizing invasive plants as sources of novel, useful traits in crops for biotechnology is our perception of what is useful, beautiful and harmful.
   
Other ecological services are provided by weeds and invasives.  Purple loosetrife (Lythrum salicaria) is a beautiful and pleasing plant species.  Zebra mussels (Dreissena polymorpha) have cleaned turbidity from many localities in the U.S. great lakes (e.g. Michigan's Saginaw Bay and Lake Huron), enhancing human health and recreational value.  Weeds are a major food source for indigenous wildlife bird and animal populations (Martin et al., 1961).  They provide refuge habitat to many other species, and can be alternate hosts for beneficial insects and microflora in biocontrol tactics.  In addition, their vast phenotypic biodiversity is valuable in its own right. 
(Dekker, 2005)
 



 
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