CENTRE FOR PLANT PROTECTION STUDIESDEPARTMENT OF AGRICULTURAL ENTOMOLOGY

TAMIL NADU AGRICULTURAL UNIVERSITY COIMBATORE, TAMIL NADU-641 003

SSNAIK, TNAU

Chairman Dr. S. Sridharan, Ph. D., Professor of Agrl. Entomology

INTRINSIC-INTER AND INTRASPECIFIC COMPETITION IN HOST – WASP PARASITOID RELATIONSHIP

DOCTORAL CREDIT SEMINAR – I (0+1)

Chairperson: Dr. S. Sridharan, Ph.D., Professor, Dept. of Agrl. Entomology, CPPS, TNAU, CBE-3.

ADVISORY COMMITTEE

Members:

ScholarS. Srinivasnaik, II Ph.DID.No: 2015800506

Dr.K.Bhuvaneswari, Ph.D.,Professor,Dept. of Agrl. Entomology,CPPS, TNAU, CBE-3.Dr.S.Mohan Kumar, Ph.D.,Professor,CPMB&B,TNAU, CBE-3.Dr.S.Nakkeeran, Ph.D.,Professor, Dept. of Plant Pathology,TNAU, CBE-3.Dr. S.K. Jalali, Ph.D., FRES Principal Scientist NBAIR, Bangalore

External

Preamble

Ways- intrinsic competition

Mechanisms in intrinsic competition

Case studies

Solitary Vs. Gregarious intrinsic competition

Fitness of progeny in intrinsic competition

PART I

PART II

PART III

INDEX

Conclusion References

Competition Strategies in intrinsic competition

Factors effecting intrinsic competition

Discussion

PART I

PREAMBLE

Integrated Pest Management Legal control Physical control Cultural control Mechanical control “Biological control” Chemical control Host plant resistance Biological control : “Parasitoids”

Predators Pathogens

Cassava mealybug-Africa Papaya mealybug-India Cottony cushiony scale- California

Parasitoid

Immature stage is parasitic Adult is free living

“Insect parasite of an arthropodan “

Adult

“Obligatory but facultative”

-Jervis, 2005

Host/prey

Phytophagy

Parasitoid/Predator

Patch

Clutch size

Phytophagous pest Parasitoid/PredatorX-Jervis, 2005

Host

Herbivore/ Phytophagy

PART II

Ways- intrinsic competition

Mechanisms in intrinsic competitionSolitary Vs. Gregarious intrinsic competition

Fitness of progeny in intrinsic competition

Competition Strategies in intrinsic competition

Factors effecting intrinsic competition

"Competition is a manifestation of the struggle for existence in which two or more organisms of the same or of different species exert a disadvantageous influence upon each other because their more or less active demands exceed the immediate supply of their common resources." Predators- Broad host range Parasitoids- Narrow host range

1. Host habitat finding 2. Host finding 3. Host acceptance 4. Host suitability

Host/Patch marking

Requisite host in less Volatility of the marking pheromone Sharing of the hosts in a patch Pro ovigeny condition

Failure of host discrimination

“Competition”

Despite of having egg, larval, egg larval parasitoids Despite of having egg, larval, egg larval parasitoids

Plant-herbivore-parasitoid systems Integration of the parasitoids-successful pest control -Paul Debach, 1973

Two types of competition: 1. Extrinsic competition 2. Intrinsic competition

Extrinsic competition Intrinsic competition

Among free-living adults Among immature parasitoids

Life cycle of a parasitoid

Egg to egg - xEgg to I instar larvae I instar to late instar

Harvey et al., 2013

Kill the host Static resources-quality

Many individuals

Single individual

Outside of the host Perforate the cuticle Imbibe the fluids Large with yolk eggs

Allow it grow the host Resources dynamic

Tissue, Haemolymph Direct contact Yolkless eggs-extra membrane Total feeder/haemolymph/fat

-Yamamoto et al.,2007 and Harvey et al., 2013

Microplitis mediator

Solitary larval endoparasitoid Gregarious larval endoparasitoid

Cotesia glomerata

Diadegma semiclausum

Solitary larval endoparasitoid

-Gols and Harvey, 2009

Two ways 1. Superparasitism 2. Multiparasitism

Superparasitism: Parasitism of a host by parasitoids of the same speciesMultiparasitism:Parasitism of a host by parasitoids of the different species

Antagonistic interactions - monopolize host resources.Parasitoids do not hesitate to oviposit in already parasitized hosts Self and Conspecific Heterospecific Superparasitism delays progeny

development of immature parasitoids Smaller parasitoid progeny Competitive displacement

-Bai and Mackauer, 1992

Two mechanisms 1. Physical defense 2. Physiological suppression

On hatching - move actively-abdomen and tail

Embeds its mandibles into the body of the other

Leakage of body fluid from the punctured cuticle(Whole of the bitten larva becomes considerably shrunken).

Larva-susceptible to the blood reactions of the host

Encapsulation by haemocytes, deposition of melanin

Distance between the members of each pair of spots distance between the mandibles

Aggregation of host phagocytes around the wound

6 days this enlarges considerably and either encloses the whole larva in a loose capsule or forms a tubular capsule around itHarvey et al., 2013

Embedding mandibles Melanin-points of wounding partial encapsulation

Melanization and a tubular haemocyte capsule

Cessation of feeding, shrinking and stunting of the larvae Encapsulation (Sub spherical/Regular outline compact capsule) Improper eclosion of the larvae

No mandible attack only contact through the haemolymph

What ?

How ?

Secretion of the toxic substance Stimulation host blood to produce more phagocytes Conditioning-haemolymph-Excretory, salivary secretions

Encapsulation

Larvae Adult I. Larvae

Harvey et al., 2013

DIFFERENTIATION ?

II. Adult Molecular tug-of-warTeratocytes, Venoms and Polydnaviruses (PDVs)

Passive and Active

Egg coating

Hatching Regulation of host

growth Synergistic effect Trophic function

-Asgari, 2006 and Harvey et al., 2013

Venom with calyx fluid Pupal stage 10-100 kda 6 major proteins

Ectoparasitoids (Venom) Endoparasitoids (Venom+PDVs)

Paralysis, Morphology, viability, and immune function of haemocytes Host immune suppression Haemocytes spreading

Indirect effect of venom Promoting PDV infection Synergistic effect

-Asgari, 2006 and Harvey et al., 2013

His research on the underlying mechanisms of insect immunity culminated in the treatise, The Cellular Defence Reactions of Insects

Placing washed eggs of the parasitoid Venturia (Nemeritis) canescens into living larvae of its host Ephestia kuehniella

Calyx proteins

Ichnovirus Bracovirus

Venturia (Nemeritis) canescensGenome-integrated wasp symbiontsSlavicek, 2012

Slavicek, 2012

Slavicek, 2012

Braco Ichno

Slavicek, 2012

Phenoloxidase enzymes convert tyrosine to dopa, and dopa to dopaquinone, leading to the formation of melanin;

Slavicek, 2012

1.Time of oviposition2.Environmental temperature 3.Host species 4.Host feed-Nasty host hypothesis

Time of ovipositionSelf/Conspecific/Hetero

Host species-The caterpillar Host feed-caterpillar feeds

-Fisher, 1961; Weil and Sagarzazu 1998 and Harvey et al., 2005

I. Oviposition at the same time Horogenes chrysostictos Nemeritis canescens

Ephestia sericarium

Methodology 15 healthy mature larvae for 4 hr

6 replicates @ 25 0C (90 Larvae)

4 DAO-dissections-Observations

-Fisher, 1961

41 of the Ephestia containing both species 36-only one larva 24 –Horogenes; 12-Nemertis

Emergence time :Horogenes : 59 + 2 hr; Nemertis : 69 + 2 hr

Changed the experimentInjection method

6 Days after injection observations

8 hosts unsuccessful16 died before conclusion of the exp. 23/18

Ratio: 1: 1 Active & undamaged Bitten Melanin deposition

ii) Oviposition at different times

-Fisher, 1961

-Fisher, 1961

The effect of temperature on development of the eggs and I-instar larvaeEgg

Larva

Endoparasitoids ecological niches-host insects Ephestia sericarium Ephestia elutella Ephestia cautella “Galleria mellonella” Achroia grisella

Horogenes chrysostictos Nemeritis canescens Development is slower

11 days- I instar moult ( 6 days in Ephestia) 32-33 days delayed emergence (22 days ) Horogenes did not effect the development

Herbivore

Parasitoid foraging

“HIPVs”Host may seek different food plants Quality of food plants is important Nutrients and the toxins - Performance of the host

“Direct indirect effects”

Two types of interactions

Plant-Herbivore-Parasitoid Parasitoid-Herbivore-Plant

koinobiont endoparasitoids Network of interactions Asymmetric fashion

Phytotoxins adaptations - influence intrinsic comp.Stronger competitor in preferred than additional. Compramisation of immune response

- Gols and Harvey 2009 and Harvey et al., 2013

“Safe host/heaven hypothesis”

Solitary parasitoids Strong mandibles in the first instar larvae. Well developed caudal appendage/tail

Without any mandibles Lack of caudal appendage/tail Larvae occupy much more confined spaces Without necessarily antagonistic encounters Scramble competition

Gregarious parasitoids

Par

asito

id lo

ad

Parasitoid fitness

- Harvey et al., 2013

Antagonistic interactions Winning competitor-Elimination of superneumeries

Effect of fitness Survival – Superparasitism/Multiparasitism/Simple Adult size-Decreased Developmental time increased Adult longevity decreased

Idiobionts-Per capita resources are lessKoinobionts-Regulate the host feeding rate and growth

-Strand et al., 1988 and Harvey, 2005 and Harvey et al., 2013

PART III Case studies

Conclusion ReferencesDiscussion

1

Microplitis croceipus Cardiochilus nigriceps Campoletis sonorensis Chelonus insularis

“Multi-interspecific-endoparasitic competition”

Methodology

Tobacco budworm

Glass container

Paper towel

Cut squares (20-30 eggs)

3/4 females4 hours

Long vial Artificial diet

Petri dish

Larvae Second stage larvae

Expose to the individual larval endoparasitoid

Emergence

Emergence of larval parasitoids

Results and Discussion Fate of larval endoparasitoids

Host discrimination M. Croceipus; C. sonorensis Physical combat No advantage

Larger size Mobility Poor mandibles

……

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……

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Cardiochilus is a poor competitor

2

Cotesia flavipes Cotesia sesamiae “Multi-Intra/interspecific-endoparasitic competition” Chilo partellus

Sesamia calamistis

3

“Multi-Intra/interspecific-endoparasitic competition”

Methodology

Collection-Mythimna separata and parasitoids

Artificial diet, INSECTA-LFS-16:8 hrs. Photoperiod, 25 ±1 o C

Mating and maturation-oviposition activity @ room temp., 30 % sugar solution

100 M. separata larvae parasitisation individually

Reared in the plastic boxes

Dissection-HEPES buffered saline solution

Observations: oviposition to cocoon formation

Morphological observations-SEM

Pair wise contest (1, 24, 48 and 96 hrs intervals

Important component is bio control: parasitoids, predators, pathogens Parasitoids important to study to understand the parasitoid-herbivore-

plant interaction Trends like development of insecticide, thermo tolerant, Diapause,

Genetic engineering & Competition Host requisite in less, provigeny condition- failure of host

discrimination Two types: extrinsic and intrinsic Extrinsic: free living adults and intrinsic: immature stages Idiobionts, Koinobionts, Ecto, Endo, Solitary and Gregarious Competition: between the egg and I instar and I instar and later instars Super and multi parasitism –Ways of competition Mechanisms: physical and physiological Physical by the mandibles and physiological by larva and adult Larvae: toxic substances, stimulation and conditioning of the

haemolymph Adult: Teratocytes, Venom, PDVs

Factors: Time of Oviposition, Temperature, Host and host feed (Nasty and Safe heaven hypothesis)

Based on the competition the fitness of the progeny would vary. Winning parasitoid fitness lesser than the simple parasitic progeny Optimum load crossed the scramble competition takes place in the

gregarious parasitoids In solitary parasitoids I instars use mainly the sickle shaped mandibles

for defense In gregarious parasitoids mandibles not well developed and dependent

on the physiological suppression Later instars also dependent on the physiological suppression

“As the parasitoids are involved in the competition because of their narrow host range. Biocontrol programmes should concentrate on the best integration methods involved with egg, larval and pupal parasitoids and their time of release. Molecular techniques should develop to identify the multiparasitic hosts”.

A VIDEO ON INTRINSIC COMPETITION

Vinson, S. B., J. R. Ables. 1980. Interspecific competition among endoparasitoids of tobacco budworm larvae (Lepidoptera: Noctuidae). Entomophaga. 25(4): 357-362.

Harvey, J.A., E.H. Poelman and T. Tanaka. 2013. Intrinsic inter and intraspecific competition in parasitoid Wasps. Annu. Rev. Entomol. 58: 333-351.

Sallam, M.N., A. William, Overholt and E. Kairu. 2002. Intraspecific and interspecific competition between Cotesia flavipes and Cotesia sesamiae (Hymenoptera: Braconidae), gregarious larval endoparasitoids of lepidopteran stem borers. Bioc. Sci. and Tech. 12: 493-506.

Magdaraog, P.M., J.A. Harvey, T. Tanaka and R. Gols. 2012. Intrinsic competition among solitary and gregarious endoparasitoid wasps and the phenomenon of resource sharing. Ecological Entomology, 37: 65-74.

Gols, R. and J.A. Harvey.2009. Plant-mediated effects in the Brassicaceae on the performance and behaviour of parasitoids. Phytochem Rev., 8:187-206.

Fisher. R.C. 1961. A study in insect multiparasitism II. The mechanism and control of competition for possession of the host J. Exp. Biol., 38: 605-628.

Asgari, S. 2006. Venom proteins from polydnavirus-producing endoparasitoids: Their role in host-parasite interactions. Archives of Insect Biochemistry and Physiology, 61:146-156.

Mayhew, P.J. and J.J.M. Van Alphen.1999. Gregarious development in alysiine parasitoids evolved through a reduction in larval aggression. Animal Behaviour, 58: 131-141.

Harvey, J.A., F. Pashalidou, R.Soler and T. Martijn Bezemer.2011. Intrinsic competition between two secondary hyperparasitoids results in temporal trophic switch, Oikos, 120: 226-233.

EC. Lampert, LA. Dyer and M. D. Bowers. 2010. Caterpillar Chemical Defense and Parasitoid Success: Cotesia congregata parasitism of Ceratomia catalpa. J. Chem. Ecol., 36: 992-998.

What is the distance between two mandibles if the distance between the two melanin spots on the body of that larva is 250 µm ? A)250 µmB)500 µmC)1200 µmD)1500 µm

Answer ?