3/11/2013  Out of the Wilds and Into Your Garden                                        Warfare in the Garden             ...
3/11/2013But plants aren’t merely generous benefactors                                                                    ...
3/11/2013          Phytochemicals: the basis of plant chemical warfare                                                    ...
3/11/2013                                                                                 S. CA Walnut: not for           ...
3/11/2013       Juglone is an effective toxin because it affects                                                          ...
3/11/2013                     Other native trees/large shrubs with                           allelopathic properties      ...
3/11/2013     Southern Goldenrod – Solidago spectabilis var. confinis                                                  Oth...
3/11/2013                The genus Solidago: the Goldenrods                                     Outside of their native ra...
3/11/2013              Why are U.S. Goldenrods so invasive in                                                             ...
3/11/2013                                                                                                                 ...
3/11/2013             What types of invaders/competitors do                                                               ...
3/11/2013    Insects are known to be selective in their                                                             Certai...
3/11/2013                      Diterpenes: another class of bioactive                                                     ...
3/11/2013                                                                                                                 ...
3/11/2013         Labdane terpinoids: typical preformed                                                                   ...
Garden warfare   2013-handouts
Garden warfare   2013-handouts
Garden warfare   2013-handouts
Garden warfare   2013-handouts
Garden warfare   2013-handouts
Garden warfare   2013-handouts
Garden warfare   2013-handouts
Garden warfare   2013-handouts
Garden warfare   2013-handouts
Garden warfare   2013-handouts
Garden warfare   2013-handouts
Garden warfare   2013-handouts
Garden warfare   2013-handouts
Garden warfare   2013-handouts
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Garden warfare 2013-handouts

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This lecture was given in March, 2013 as part of the California native plant gardening series ‘Out of the Wilds and Into Your Garden’

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Garden warfare 2013-handouts

  1. 1. 3/11/2013 Out of the Wilds and Into Your Garden Warfare in the Garden how plants protect themselves from pests and invasions C.M. Vadheim and T. Drake CSUDH & Madrona Marsh Preserve Madrona Marsh PreserveGardening with California Native Plants in Western L.A. County March 2 & 5, 2013 Project SOUND – 2013 (our 9th year) © Project SOUND © Project SOUND Plants are primary producers Why consider plant defenses?  Plants – and everything about them – are inherently interesting  Understanding how plants interact with other living things may improve your gardening  Plants and animals are more similar than we think – at least at the cellular level Photosynthesis: stored energy,  May suggest novel medicines, pesticides and other useful other (biomass) prodcucts http://www.bostonbakesforbreastcancer.org/summer-sun-radiation-and-chemo/ http://www.glogster.com/beckeyy/food-web/g-6mp96eehhgdfvco22h8bna0  That means they are ‘food’ to many organisms © Project SOUND © Project SOUND 1
  2. 2. 3/11/2013But plants aren’t merely generous benefactors After all, plants have to live too… or faithful servants….  Capturing energy needed for life (photosynthesis)  Obtaining water: roots  Obtaining nutrient chemicals (primarily through the roots)  Reproducing: seed or vegetative  Protecting themselves from anything that impacts the above:  Abiotic factors: temperature, weather, soils etc.  Biotic factors: living things http://thebillfold.com/2012/04/reader-mail-how-to-be-a-generous-person/ CA Goldenrod - Solidago californica © Project SOUND © Project SOUND Plants are far more Types of defenses complex than most  Physical barriers (preformed or of us realize induced)  Waxy cuticle  They interact with many types of  Trichomes/leaf hairs organisms – for better or worse  Tough epidermis  They are at least as complex as  Sticky gums & resins higher animals  Prickles, barbs and thorns  Dense growth habit  They were not set on this planet  Hard covering to protect seeds for our use alone http://img.ehowcdn.com/article- new/ehow/images/a08/4f/it/avoid-leggy-seedling- 800x800.jpg  Etc.  They often behave more like  Defense chemicals plant warriors than like shrinking  Preformed: always ready violets  Induced: produced only when needed (usually when stimulated by an attack) © Project SOUND © Project SOUND http://gorillaartfare.com/character-design/two-little-kittens/ 2
  3. 3. 3/11/2013 Phytochemicals: the basis of plant chemical warfare Phytochemicals  The distribution of phytochemicals within plants is often tissue/organ specific  Phytochemicals: non-nutritive  These molecules tend to be concentrated plant metabolites that are in outer cell layers of plant organs, essential for the survival and suggesting that they may indeed act as proper functioning of growth and deterrents to pathogens and pests. reproduction in plants  These compounds are of two types:  Often involved in protection  Constitutive chemicals: made during against herbivores, pests and normal growth and development micro-organisms (or other (preformed antimicrobial compounds, or environmental stresses) “phytoanticipins”)  Sometimes used by animals that http://plantpathology.uark.edu/4844.htm  Induced chemicals: absent from healthy eat them: plants, accumulating only in response to pathogen attack or stress (‘phytoalexins’)http://naturallyjodi.blogspot.com/2012/08/medical-news-  As defense chemicals2012.html#!/2012/08/medical-news-2012.html  As human plant-based medicines,Phytochemicals are sometimes flavorings and other uses Whether a given compound has a defensive function is thecalled secondary metabolites subject of much current interest & research © Project SOUND © Project SOUND Competition between plants: it’s a fact of life, Southern CA Walnut – Juglans californica particularly in our mediterranean climate  Light  Water  Soil nutrients  Access to services of beneficial organisms Plants sometimes have to ‘fight dirty’ to best their competitors: allelopathy http://primarybestsc.blogspot.com/2012/10/plants-competition.html http://the-gist.org/2012/09/allelopathy-when-plants-attack/ http://biology.csusb.edu/PlantGuideFolder/JuglansCalif/JuglansCalifPage.htm © Project SOUND © Project SOUND 3
  4. 4. 3/11/2013 S. CA Walnut: not for Allelopathy: one type of chemical warfare every yard  Term from allelon ("of each other“) and pathos ("to suffer“).  Proper location: consider  Refers to the chemical inhibition of one species by another.  Size: moderate for tree  Allelopathy has generally come to mean the deleterious effect  Light: full sun/part shade http://biology.csusb.edu/PlantGuideFolder/JuglansCalif/JuglansCalifPage.htm that one plant has on another through the production of  Soils: clay soils best chemical retardants  Water regime:  The "inhibitory" chemical is released into the environment where it  Tolerates seasonal flooding affects development and growth of neighboring plants.  No or very occasional deep water in summer (hot gardens)  Process is often more complex:  Allelopathy:  Allelopathic plants are also capable of stimulatory effects  Leaves produce chemicals toxic  The chemical producing plant may also inhibit itself with the same to other plants chemicals that inhibit its neighbors  Can’t grow plants under walnuts  The process may involve other organisms [soil microbes]http://www.phytoimages.siu.edu/imgs/paraman1/r/Juglandaceae_Juglans_nigra_4178.html © Project SOUND © Project SOUND Plants release allelochemicals in several Naphthoquinones different ways  Often responsible for colored barks, root barks and heartwoods http://kremerpigments.com/shopus/index.php?cat=0104&lang=EN  Example: Juglone (C.I. Natural Brown 7) G&product=37300&sidFEE4B14F27014E7795A5F1BD0DD62743=  Above ground 63e5300d98a56f6479a23d579380ca6a  Leaves release volatiles  In leaves, roots, husks, and bark of plants in the Juglandaceae family  Leaching from leaves  Is toxic or growth-stunting to many types of  Leaching from plant litter or plants and insect herbivores - inhibits key on decomposition enzymes needed for metabolic function.  Below ground  Awareness of walnut toxicity dates back at least to Roman times  From above-ground leachates  Used as:  Root exudates  an herbicide  Decomposing roots  a dye for cloth and inks  a coloring agent for foods and cosmetics (hair dyes).  Folk medicine – ground/extract green hulls http://pubs.ext.vt.edu/430/430-021/430-021.html © Project SOUND © Project SOUND 4
  5. 5. 3/11/2013 Juglone is an effective toxin because it affects The toxin is stored in a non-toxic form (so basic processes required for life it doesn’t harm the walnut tree)  Juglone exists within wlanut tree cells in a  The active agent inhibiting growth of non-toxic form called hydrojuglone. other plants was suggested by Massey in 1925; confirmed by Davis in 1928.  Hydrojuglone is colorless and generally nontoxic, but is immediately converted to  Juglone disrupts oxygen and food use in juglone by oxidation. both plants and animals, a respiration http://www.biologie.uni-hamburg.de/b-online/ge20/02b.gif  Upon continual contact with oxidative poison. Juglone is like cyanide in its conditions, or tissue drying, juglone is tiedhttp://www.bgshoppingmall.com/Brand-Names/abbyson-living.php effect on people, animals, and plants up and decomposed.  Juglone is so toxic only minute amounts  When you cut open a green walnut husk, it can sicken, sedate, or kill people and quickly turns brown when exposed to air. So how do walnut animals. The concentration difference This is caused by the clear, non-toxic trees survive? of juglone between that needed for hydrojuglone being quickly converted into sedation, and that causing death, is the toxic, dark brown juglone in the small. presence of oxygen. © Project SOUND © Project SOUND Why do Juglans species make juglone? At Allelochemicals: many modes of action least part of the story involves seedlings  Allelopathic chemicals can be  Juglone in the husk protect the seed from being present in any part of the plant - eaten. The juglone also leaches into the leaves, flowers, roots, fruits, or surrounding soil stems.  If juglone leaks back into a walnut root, it is  They can also be found in the quickly made non-toxic again and stored. surrounding soil.  Annual plants, garden vegetables, fruit trees, and some broad-leaf perennials can be severely  Target species are affected by damaged when juglone is in the soil. These are a these toxins in many different ways: seedling’s biggest competitors  Inhibited shoot/root growth http://www.hiltonpond.org/images/WalnutBlackS  Most grasses seem immune from juglone  Inhibited nutrient uptake eedling01.jpg problems.  Altered symbiotic relationship [mycorrhyzae] - destroying the  Select mycorrhizal fungi and soil microbes have plants usable source of a nutrient. been shown to be highly adapted to walnut tree control zones and the presence of juglone. http://www.sustland.umn.edu/implement/images/trees_turf_4.gif © Project SOUND © Project SOUND 5
  6. 6. 3/11/2013 Other native trees/large shrubs with allelopathic properties Dealing with allelopathy in the garden  Rake up leaves & other parts containing the chemicals(s)  Oaks – Quercus spp.  Leaves, twigs, fruit husks, and wood chips from walnut trees  Cottonwoods – Populus spp. should be well aged or completely composted before adding  Manzanitas – Arctostaphylos spp. to a garden or landscape, if at all  CA Sycamore – Platanus racemosa  Walnut stumps should be ground-down or removed from a  CA Bay Laurel - Umbellularia site californica  Plant tolerant species under/near: http://farm1.static.flickr.com/29/101896704_625b8ccece.jpg  False Indigo - Amorpha fruticosa  Solanaceae, annuals are particularly susceptible  Grasses are usually not  Eucalyptus  Soil microorganisms ingest allelochemicals as energy sources,  Tree of Heaven and metabolic decomposition can render the chemicals non- toxic to plants. When soils are well drained and aerated, a healthy population of aerobic microorganisms can accelerate this progress. © Project SOUND © Project SOUND But making Juglone is expensive: is Pros/cons of preformed defense allelopathy the only explantion? chemicals (or precursor forms of them)  Juglone precursors appear to be  Positive translocated from older tissue to younger tissue over time.  Always ready  Mechanism often involves basic  The immediate precursors of mechanisms – effective against a juglone are found in high wide range of living things concentrations within buds, flowers, fruit, and in the phloem http://enhancedbc.tfrec.wsu.edu/CA_walnut.html (vascular system).  Negative  May not ever be needed – a waste of  Juglone is also effective for energy & other resources protection from leaf, root and  Not specific – so may not work as well stem pests, like insects, diseases, against some threats nematodes, and grazing animals.  May be deleterious to the plant itself http://c0365781.cdn2.cloudfiles.rackspacecloud.com/datas/5598254/http://www.graftedwalnuts.co.uk/pest.ihtml original/799px-155mmMustardGasShells.jpg © Project SOUND © Project SOUND 6
  7. 7. 3/11/2013 Southern Goldenrod – Solidago spectabilis var. confinis Other good native Goldenrods http://www.jcsemple.uwaterloo.ca/goldenrod_figs.htm Solidago californica Euthamia (Solidago) occidentalis © Project SOUND © Project SOUND The genus Solidago: the Goldenrods Southern Goldenrod is an herbaceous perennial  ~ 100 perennial species  Size:  2-3 ft tall  Most grow in meadows, pastures, along  2+ ft wide, spreading roads & ditches in North America  Unfairly blamed for hay fever in late  Growth form: summer/fall - Ragweed (Ambrosia sp.),  Stout looking herbaceous blooming at the same time but wind- perennial pollinated, is the usual culprit.  Fall/winter deciduous; dies back to basal rosette  Easily recognized by their golden flowering stalks with hundreds of small  Foliage: flowers; plants & flowers make nice  Leaves lance-shaped – mostly yellow & green dyes. basal  Their alternate leaves are linear to  Leaves fleshy, bright to paleCA Goldenrod - Solidago californica lanceolate. Their margins are usually green finely to sharply serrated.Goldenrods have been used in  Roots: spreads via rhizomesBritish gardens for > 200 years © Project SOUND © Project SOUND © 2003 Christopher L. Christie 7
  8. 8. 3/11/2013 The genus Solidago: the Goldenrods Outside of their native range, Goldenrods  Propagation by wind- can be invasive. Why? disseminated seed or underground rhizomes (form patches that are vegetative clones of a single plant).  Goldenrod is a companion plant,CA Goldenrod - Solidago californica playing host to beneficial insects, repelling some pests  Goldenrods are important habitat plants for a wide range of native insects, butterflies, birds, etc. © Project SOUND © Project SOUND http://www.calflora.net/bloomingplants/southerngoldenrod.html http://www.calflora.net/bloomingplants/southerngoldenrod.htmlTeasing out whether the effect is due to allelopathy: Bioassays often used to test for allelopathycan be difficult  Competing processes: competition for  Light  Water  Nutrients  Associated animal species may be the culprit:  Pollinators, mycorrhyzae, other beneficial species  Above or below-ground pests – Solidago http://www.biosci.ohio-state.edu/~plantbio/osu_pcmb/pcmb_lab_resources/images/pcmb300lamb/allelopathyExperiment.jpg may be tolerant  Example: testing the effects of plant tissue extracts (or specific  Vast number of chemicals produced; chemicals) on the germination of seeds. many not toxic (at least to other plants)  Issues in relating laboratory bioassays to allelopathic  Nature of the chemicals themselves: interactions in the field; allelopathy in the laboratory is not highly changeable (oxygen; pH; exposure always demonstrated in the field – and vice versa to other chemicals) http://plantecology.dbs.umt.edu/People/collaborators.html © 2003 Christopher L. Christie © Project SOUND © Project SOUND 8
  9. 9. 3/11/2013 Why are U.S. Goldenrods so invasive in The answer is not exactly straightforward China? Could it be due to allelpathy?  Solidgo canadensis does influence soil levels of possible allelochemicals (total phenolics, total flavones and  Extracts were made from 12.5 g of dried total saponins) leaf tissue placed in 500 ml of distilled water.  The chemical content and possible allelopathic effects were greater in  Dilutions of each extract, ranging from S. canadensis from China than 0% to 100% in 10% increments were those from the USA as demonstrated made. in a field survey and a common garden experiment.  Filter paper was placed in 90 mm petri http://www.sciencedirect.com/science/article/pii/S0929139311000849 plates with 20 seeds of the target  Suggests that S. canadensis has species (lettuce & radish). But is the effect direct? evolved to be more competitive – and possibly more allelopathic - in the  Five trials were run for each dilution for introduced range each goldenrod species tested. http://posieinthevase.blogspot.com/  Allelopathy might significantly increase competitiveness for this © Project SOUND invasive species. © Project SOUND Plants have complex relationships with Roots and the rhizosphere: life within the soil other living things Beneficial effect of mycorrhyzaehttp://www.sciencedirect.com/science/article/pii/S1360138510001007 http://ars.els-cdn.com/content/image/1-s2.0-S1360138512000799-gr2.jpg © Project SOUND © Project SOUND http://www.cottoncrc.org.au/industry/Tools/Symptoms_Identification_Tool/Cotton_Symptoms/Allelopathy 9
  10. 10. 3/11/2013 A novel, non-native plant species (like Goldenrod) Goldenrods brought ‘novel weapons’ with them can have many effects on the environment  S. canadensis altered the indigenous mycorrhizal fungal spore composition and reduced the mycorrhizal colonization of native plants one year after invasion.  The alien Goldenrod inhibited the colonization of native species and changed the indigenous mycorrhizal fungi by exuding allelochemicals.  Experimental results suggest that http://openi.nlm.nih.gov/detailedresult.php?img=2972720_pone.0015418.g001&query=the &fields=all&favor=none&it=none&sub=none&uniq=0&sp=none&req=4&simCollection=305 invasive S. canadensis may acquire 8081_1471-5945-11-5-6&npos=74&prt=3 spreading advantage in non-native habitat by using “novel weapons” to inhibit not only local plants but also soilborne pathogens and beneficial microbes. © Project SOUND http://www.sciencedirect.com/science/article/pii/S016953471000145X © Project SOUND You may have noticed that the Sunflower family is pretty successful Possible reasons for Sunflower success  The Sunflower family  Ability to adapt well to variety (Asteraceae) is one of the of environments most diverse families in  Plasticity: changes in phenotype California.  Ability to evolve quickly – change in the genes in the population  Largest plant family worldwide: contains ~ 1550  Make lots of seeds genera and 24,000 species.  Work well with wide range of  Almost 200 pages of the pollinators and other beneficial Jepson Manual are dedicated specieshttp://www.wildflowers-and-weeds.com/Plant_Families/Asteraceae_pics/Asteraceae.jpg to describing the California  ?? Good defenses species alone. Why are they so successful? © Project SOUND © Project SOUND 10
  11. 11. 3/11/2013 What types of invaders/competitors do Young leaves and other tissues are plants need to worry about? attractive food  Other plants (compete for water, light, nutrients)  Large herbivores  Insect herbivores  Other herbivores: mollusks  Pathogens  Fungalhttp://www.sciencedirect.com/science/article/pii/S1360138509003008  Bacterial  Viral © Project SOUND © Project SOUND Why do plants make the chemicals found The ‘Double-whammy’ of injury in ‘essential oils’? http://ericwongmma.com/wp- content/uploads/2013/01/double- whammy.jpg  Tissue damage (from physical  A wide range of reasons, many injury or herbivory) related to communication:  Vascular tissue: must seal off quickly, then re-grow or re-attach  To attract pollinators – or the if possible spreaders of seed (usually in  Water – Xylem tissue flowers, fruits)  Nutrients + other - Phloem tissue  To repel herbivores – insect or  Support tissue other; either cue or toxin  Other http://www.tantebazar.com/gardening_histology_of_plant_part_2.php  As breakdown products from compounds used for other purposes  Secondary infection  As protection against fungi, bacteria  Bacterial and viruses  Fungal  To prevent other plants from  Viral growing too close ?  To communicate with other plants – via soil water or air © Project SOUND http://ipm.ncsu.edu/corn/diseases/cornfg18.gif © Project SOUND 11
  12. 12. 3/11/2013 Insects are known to be selective in their Certain plant compounds are effective use of plants in the sunflower family against insect herbivores http://ipmworld.umn.edu/chapters/charlet2.htm Long-horned Beetle http://michellebiology.blogspot.com/2012/02/54-pesticide-and-biological-control.html Sunflower Bud Moth  The most important of these are alkaloids, terpenoids, steroids, phenols, saponins and tanninsThis suggests that Asteraceae may selectively deter some pest species  These may be an alternative source of insect control agents © Project SOUND © Project SOUND The most cost effective strategy is often Mint family insecticidals: mostly terpenoids to prevent/limit herbivory  Most common : monoterpenes (1,8-cineole, thujone, camphor, pulegone, menthone, others)  Has a pleasant odor similar to pennyroyal, peppermint and  Plant-derived insecticides may represent camphor. alternative pest control strategies.  They may degrade more rapidly than the  Is used in flavoring agents, in synthetic insecticides perfumery, and in  May be more specific in their action aromatherapy  Have no genotoxicity. Pulegone  Mint oil is already used as an environmentally-  Insect repellant; less toxic to friendly insecticide for some common pests like animals/humans than other wasps, hornets, ants and cockroachesPlants in both the Sunflower insect repellantsand Mint families make a range  Mints also repel some birds & other largeof chemicals to prevent/limit herbivores – terpenoid’s smells repelherbivory © Project SOUND © Project SOUND http://www.safesolutionsinc.com/TweetMint_Gallon.jpg 12
  13. 13. 3/11/2013 Diterpenes: another class of bioactive How do plant insecticidal chemicals work? terpenoids  Feeding deterrants:  They have 20 carbon atoms  Render plants unattractive or unpalatable;  Produced by plants and fungi; often play  Usually small chemicals; may be aromatic active role as hormones (Gibberelins)  Often bitter or strong tasting/smelling  Examples: alkeloids, terpenopids;  Found in resins, gummy exudates, and in the resinous high-boiling fractions  Direct toxicity: remaining after distillation of essential http://ars.els-cdn.com/content/image/1-s2.0- S0367326X02001703-gr1.gif http://www.gov.mb.ca/agriculture/crops/ins  Kill insects outright; or stun them ects/images/fae02s00a.jpg oils. significantly so that they are eaten by their  Diterpenes display a broad range of predators activities against insects  Usually function as neurotoxins  Examples:  Important defense chemicals in Asteraceae, Salvia, many others monoterpenes  Other, more subtle methods: © Project SOUND © Project SOUNDhttp://www.cyberlipid.org/images/pict295.gif How do plant insecticidal chemicals work? California/Big Gum Plant A few more subtle means Grindelia camporum var. bracteosa  Modifying plant food absorption  Modify either the food itself, the gut wall or gut flora  Often larger size  Example: Tannins – make food undigestable  Disrupting the endocrinologic balance of insects  Affects reproduction  Acting as insect growth regulators, disrupting the normal process of morphogenesis  May ultimately kill  Usually affects reproduction  Behaviour modifying agents  Usually influence the feeding and ovipositing (egg-laying) behavior of insects © Project SOUND 13
  14. 14. 3/11/2013 Asteraceae species do much to protect CA Gum Plant their young leaves & flowers  Erect herbaceous perennial to 4 ft tall by 4 ft wide  Grows in clay or sandy soil:  Dry stream banks, washes  Rocky fields & plains  Sandy or alkali bottomlands  Along road sides  Grows where it gets full sun  Is stress deciduous – looses leaves during dry periods © Project SOUNDGumplants make an interesting array of chemicals Protecting developing leaves and flowers  Grindelia species are used as food plants  Active compounds: resin flavanoids by the larvae of some Lepidoptera species and diterpenes of the grindelane type. including Flower Moths, other herbivores  The resin produced in multicellular  Grindelane diterpenoids make up most of glands on the surfaces of stems, the resin (to 20% of the dry weight). leaves, and involucres - density of  Grindelic acid, camporic acid, resin glands highest on the immature  17-carboxygrindelic acid involucre bracts and lowest on the  many other diterpenoids stems.  The diterpenoids appear to have insect  The resin is composed of grindelic repellant/insecticidal & antibiotic activity acid and several of its derivatives. These labdane diterpenes are similar  Used as a traditional medicine (until 1960) to the resin acids that constitute  wide range of ailments: asthma, bronchitis; Balsamic scent – fairly strong rosin, a principal product of the naval antispasmodic , urinary tract disinfectant; topical preparations to soothe burns, insect stores industry bites, skin rashes, poison ivy rash. © Project SOUND © Project SOUNDhttp://www.ag.arizona.edu/~spmcl/Research/newcrops.htm 14
  15. 15. 3/11/2013 Labdane terpinoids: typical preformed Have you ever noticed how birds know compounds affecting a wide range of pests when the fruits are ripe?  Stored where likely to be needed  Resin ducts of trichomes of leaves  In special plastids in epidermal or other near- surface cells  A variety of biological activities:  Antibacterial, antifungal, antiprotozoalFound in many plant  Anti-inflammatory activities and modulation ofparts (above/below- immune cell functions – may induce long-termground) immunity  Significant effects on basic cell processes:  Interfere with biochemical pathways of cell death and the cell cycle phases  May explain why they affect wide range of cell types in pests (insects, microbes) and in humans http://www.gardenguides.com/633-barren-bushes-treat- hungry-birds.html © Project SOUND © Project SOUND Many fruit-berry plants use critters to Toyon/California Christmas Berry – spread their seeds Heteromeles arbutifolia  Berries attract the critter  Critter eats the fruit –yum!  Critter moves around while seeds pass through the digestive tract  Seed are deposited – with a load of fertilizer – in a new place away from the parent plant Cedar Waxwing gobbling up Toyon fruits  How do the birds know the fruits are ripe?http://www.ibabuzz.com/garybogue/2009/04/15/cedar-waxwings-they-will-eat-no-toyon-berry-before-its-time/ © Project SOUND 15

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