PARENTAL CARE IN INSECTs
Archana Anokhe
Roll. No - 20416
INTRODUCTION
THE EVOLUTION OF PARENTAL CARE IN INSECTS
FORMS OF PARENTAL CARE
MALE VERSES FEMALE CARE
HORMONAL REGULATION & CHEMICAL SIGNALING IN PARENTAL CARE
COST& BENEFITS OF PARENTAL CARE
CASE STUDY
INTRODUCTION
Parental care - any post-ovipositional behaviour that promotes survival, growth, &
development of offspring
(Tallamy et al.,1984)
A prime example - altruistic trait evolved to enhance the fitness of the recipients
(offspring) at the expense to the donor (parents)
(Royle et al., 2012)
Classified into 3 groups:
Protect offspring
Protect resources needed by the offspring
Improve resources
(Wood et al.,1991)
THE EVOLUTION OF PARENTAL CARE (PC) IN
INSECTS
Ecological factors - harsh environments, ephemeral food sources or predation
pressure
Harsh
environment
Ephemeral
food
Predation
pressure
Contd..
Harsh environmental conditions
Egg attendance provides an alternative route to resist harsh environment
For e.g.- staphylinid beetle, Bledius spectabilis live in intertidal saltmarsh, to prevent
flooding of their nest and anoxia of their eggs, females provide care in the form of closing &
reopening of the burrow
Ephemeral or distant food sources and specialized foraging
Burrower bug, Sehirus cinctus nymphs only eat seeds of Prunella spp. & are available for a
few weeks of spring, so mothers has to compete for this resource with in a short time span
Specialized food of Cryptocercus spp. underlies the evolution of PC
(Tallamy et al., 2007)
Contd..
Cont..
Do natural enemies drive the evolution of PC ?
Predation play an important role in the evolution of parental care
In the shield bug, Elasmucha spp. females shelter the eggs and nymphs by covering
them with their body and fanning their wings when attacked
European beewolf (Philanthus triangulum), females provision brood cells with
paralysed honeybee
(Tallamy et al., 2007)
FORMS OF PARENTAL CARE
Diversity of social behaviour among the invertebrates - asset and
challenge
Richness present in both number & forms of care provided by
parents
50 families of arthropod have evolved parental care
(Costa et al.,2006)
Cont..
Types
1. Trophic egg productions
2. Attending eggs and offspring
Example
Plassid beetle Cylindrocaulus patalis
Treehopper Umbonia crassicornus
3. Protection and facilitating feeding of
mobile young
a.
b.
c.
d.
e.
Guarding young
Poised to defend
Egg dumping or brood mixing
Formidable mother
Blocking entry
a.
b.
c.
d.
e.
Fungus feeding beetles Pselaphacus sps
Assassin bug & Shield bugs
European Earwig Forficula auricularia
Praying mantis Oxyophthalmellus somalicus
Myrmecine and Zacryptocerus varians
5. Nest building and burrowing
a.
Hiding evidence
b.
Deceiving enemies
a. Solitary wasp
b. Bembix sand wasps
6. Brooding behavior & viviparity
Webspinner (Partenembia reclusa)
Contd..
1. Trophic egg productions
To making the correct choice of which eggs to consume, chemical cues is incorporated into the
trophic egg
Passalid beetle ,if 3rd instar larvae stridulate, they are fed by trophic egg
(Philipi et al., 2008)
Burrower bug , Sehirus cinctus produce some trophic eggs at the time when they produce viable
eggs as well as after the nymphs hatch. It provide food , when mother go for foraging
(Hironaka et al., 2005)
Contd..
2. Attending eggs and offspring
Treehopper Umbonia crassicornus, mothers tilt their elongated pronotum and fan the potential threats,
protecting their young until adulthood
Offspring produce synchronous vibrations that inform the mother on which side threat is more
prominent
(Ramaswamy et al., 2009)
3. Protection of mobile young
Mobile young feeding in open area vulnerable to predators & parasitoids
Females standing nearby, not only threatening potential predators, but also stroking wandering
nymphs (Lace bug)
(Windsor et al., 1994)
Contd..
a.
Guarding young:- Brazilian sawfly, Themos olfersii, continues guarding their larvae even when they
fully sclerotized & starts feeding
b.
Blocking entry:- In myrmecine colony major worker blocks entrance with its saucer-shaped head &
receives regurgitated liquid from a minor worker
Matthews & Matthews, 2010
Contd..
c.
Poised to defend :- Assassin bug & Shield bugs continuously watches her emerging
nymphs to
defend her predator
d. Formidable mother :-
The praying mantis Oxyophthalmellus spp. positioned
herself at base of a
twig to intercept the predators
e. Egg dumping or brood mixing :- Spatial proximity leads to brood mixing & alloparental care
communal breeding, egg dumping (lace bug) ,take-over of nests & offspring
burying beetle
is seen in case of
(Smith et al., 2012)
Contd..
4. Nest building & Burrowing
1. Deceiving enemies :- A bombyliid bee fly, Parasitoid of solitary wasp larvae lay several
eggs into the fake nest entrance made by the wasp
2. Hiding the evidence:- Synchronous motions of the front tarsi , rake the sand & producing a
characteristic pattern which hide its actual nest eg:-Bembix sand wasps
Contd..
4. Brooding behavior & viviparity
An alternative form of protection is to carry eggs or young either internally or externally
(Kolliker et al., 2013)
Contd..
2013 The Royal Entomological Society, Ecological Entomology, 38, 123137
FEMALE VS. MALE CARE
Parental care is indication of higher evolution
Types of care provided by either of the parent vary with the insect to insects
Dermapterans guard their egg until they hatch & cockroaches carry them in protective ootheca
(Costa, 2006)
MATERNAL CARE
o Maternal care is most exclusive form of parental care
o Damsel bug is a fearsome enemy of Lace bug , if given a chance will devour every last
nymph ,such case she diverts enemy by fanning her wings and climbing on its back
o Also guide the nymphs to an ideal leaf by blocking a branch which they mistakenly follow,
all too often she dies in the attack
(Noldus et al., 1991)
Contd..
Tortoise beetle Acromis sparsa arrange themselves into a symmetrical ring around the
hatchlings & allow them to reach near food sources in round up manner
Harlequin stink bug Tectocoris spp. defends her eggs aggressively because she lays only one
batch & are her sole chance for reproductive success
(Ken Preston et al., 2010 )
Cont..
P physical (nest, burrow, gallery tunnel, brood ball, etc.); B behavioural (i.e. defence
display, guarding); C -chemical (defensive pheromone, aggregation pheromone, alarm
pheromone , antimicrobial substance )
(Goershon et al., 2001)
Contd..
Many insects raise their offspring on organic substrates where Microbes is serious threat to offspring
development and survival either by decomposing their food resources or by directly infecting the offspring
Objective: To test the maternal egg attendance in the European earwig has a function as a social defense against mold
infection
To see the effect of egg attendance on hatching success
To investigated the potential roles of egg grooming behavior and maternal transfer of chemicals on hatching
Contd..
Result & Discussion
Experiment 1: - Effects of maternal egg attendance with and without mold exposure on hatching success (A)
and mean hatchling body weight (B)
Effect of mold spore exposure on the duration of maternal egg grooming was made over the course of a 75-min
observation period
Control
Treated
B
Contd..
Parameter
Control
Egg attendance
Hatching success (Chi square)
Hatching success (%)
Body weight (Mg)
Treated
not attended
Egg attendance
138.08
130.21
59.85
80
50
75
1.68
1.78
1.57
not attended
55.34
7
1.98
Contd..
Experiment 2:- Effect of maternal presence on chemical compounds on egg surface
Change of total hydrocarbon quantities over time & total hydrocarbon quantities on
glass beads mingled into maternally attended clutches and unattended clutches
The major Chemical were identified as
heneicosane (C21H44), Z-9-tricosene
(C23H46), tricosane (C23H48), Z-12pentacosene (C25H50) & internal standard
n-octadecane (C18H38)
Contd..
1. Maternal egg attendance reduced the detrimental effects of spore exposure
2. Spore exposure significantly increased the duration of maternal grooming
3. Attendance lead to incorporation of chemical compounds (hydrocarbons)
on the eggs
PATERNAL CARE
Exclusive paternal care has evolved in 8 order of insects
Males can improve females mating success by providing pre-zygotic nuptial gifts & postzygotic offspring care
Two ways in which female choice promote the expression of arthropod paternal care
a) Enhanced Fecundity Hypothesis
b) Handicap Principle
(Tallamy et al., 2001)
Contd..
By M. Kolliker et.al., 2000
Although sexual selection and natural selection can favour paternal behaviour, a number of parameter can help to
determine which has dominated in due course of evolution
Trait
Sexual selection
Natural selection
Clutch number produced
Iteroparous (>1)
Semeplarous (1)
Mating opportunities
Many
Few
Care versus foraging
Care interferes with female foraging Care may or may not interfere with
female foraging
Female preference
Preference for guarding male
No preference for guarding male
Clutches guarded simultaneously
>1
Paternal selfishness
Males will guard unrelated eggs
Males will not guard unrelated eggs
Female
Male
Sex reversal role
After mating, receptive males perform
vigorous pumping
Females respond by climbing on the
males back
The male abruptly scrubs the females
hemelytra,
female repositions herself
and begins to lay eggs
After 5 minute, male temporary forcing
the female out of & begins a new bout of
vigorous pumping
This cycle continues until the females
total egg clutch is deposited on the
males back
Case study
Reversed sex roles in the giant water bug, Abedus sps.
Female water bugs abandon their eggs to males that proceed to care for them. What has caused
such an unusual behavior?
An egg-encumbered male is placed in an aquarium, highest number (97%) of the eggs will hatch
Does this high egg survival rate depend upon eggs being carried on the back of a live male water
bug? If so, why?
Removing egg pads from the backs of six males, Smith placed each egg pad in a separate
fingerbowl in the laboratory ,none of the eggs were survived
Was something about the male exoskeleton necessary to their development?
Smith killed egg-bearing males and left them with eggs intact , but these eggs also failed to hatch
So attachment to a healthy male essential for hatching. Why?
He noticed, male keep egg pad at or above the waters surface. Perhaps regular exposure to
atmospheric air is required for development
(Robert smith,1970)
Contd..
Reasons for paternity
Females always prefer males which were already guarding egg
of another female
Males
willing to guard more than one clutch to promote re-
mating
A male always
attractiveness
willing to guard unrelated eggs to enhance his
BIPARENTAL CARE
Dung beetle
Female attends the offspring until
they pupate
She cleans mold growth & applying
antimicrobial chemicals derived from
her sternal glands
Male helps to dig
compact the dung pat
the
nest
&
Food relocation to nests reduce the
predation risk
(Matthews & Matthews 2010)
PHYSIOLOGICAL MEASUREMENTS OF PARENTAL CARE
Mechanism of parental care By direct internal chemical signals (i.e.
hormones) and indirect external chemical signals (i.e. pheromone)
(Kolliker et al., 2007)
It influence maternal reproductive physiology and future reproduction
Contd..
Chemical signalling - important mode of communication between parent & offspring
Apis mellifera, larvae produce a brood pheromone that influences care behaviours of
workers
JH, regulate the trade-off between care for current offspring and the production of new
eggs/clutches
JH
(Mas et al., 2008)
Contd..
Hypothetical mechanism for maternal care regulation
JH
Age, Nutritional state & JH
(Scott & Panaitof 2004 )
Host manipulation for offspring care
1.
Hymenoipimesis spp
Plesiometa argyra
2.
Ichneumon eumerus
Maculina rebeli, a butterfly parasitize by wasp
(Delago et al., 2009)
Contd..
3.
Bee wolf (Egyptian digger asp) Philanthhus triangulam known as paralysis inducing wasp. Its venom
contains potent neurotoxin (Philathotoxin) that evoke neuromuscular paralysis Of Bee prey
(Delago et al., 2009)
Contd..
4.
Sphecid wasp Ampulex compressa
paracitising Periplanata americana
for egg laying
(Delago et al.,2009 )
COST & BENEFIT OF PARENTAL CARE
Parental investment benefits a current progeny at the expense of future reproductive
success
Protection of eggs is one of the most basic forms of parental care seen in many insects
(Clutton et al., 1991)
Contd.
QUANTIFYING THE COSTS AND BENEFITS OF PARENTAL CARE IN
FEMALE TREEHOPPERS
Case study
( ZINK et. al., 2003)
1. Benefit of egg guarding: increased hatching success
There is Correlation between the hatching success, offspring survival & total number of
0.0007
21.5 @ 0.23 Nymph/egg/day
No. of nymph
guarding days by female presented below
0.804
0.483
Contd..
Costs of egg guarding: decreased future reproduction
Early abandonment of eggs allowed females to increase the overall number of future broods
1.23
6.4
14.4
4.6
0.62
10.0
Means of the future number of broods initiated (A), the mean length of those future broods
(B), and the mean future guarding duration (C) for females abandoning versus remaining for
Contd..
Parameter
Control (Egg guarding female)
Treatment (Non egg guarding
female)
Hatching success (In terms of per female)
0.007 nymph/female
0.005 nymph/female
0.05 nymph /egg
0.03 nymph/egg
0.804 nymph /egg
0.483 nymph /egg
More i.e. 2.1
Less i.e. 1.02
(In terms of egg)
Survival of Hatchling
Presence of mutual friend ant
( After hatching)
Contd..
Cost
Benefits
Egg guarding represents an
investment in terms of time , energy
& reduced future fecundity
Opportunities for laying additional
clutches will be reduced
Egg guarding by adults acts to increase the
hatching success
Offspring survival in guarded clutch will
increase increase
The egg guarding in Publilia concava shown the positive linear relationship between guarding duration and
hatching success
Females that abandoned immediately after egg laying suffered a 50% reduction in hatching success
Effect of Parental Care on the Duration of Larval Development and Offspring
Survival in Nicrophorus mexicanus Matthews (Coleoptera: Silphidae)
By -Sofa et. al., 2011
Objective of study
1. To see the effect of parental presence on larval development
2.
Effect of parental manipulation of the food resources on offspring survival
3. To see the difference in number of adult emerges during different treatment
Result & Discussion
Differences in time development & number of adult emergence among the three experimental groups in N.
mexicanus 1.Control group; 2.Brood-mass present and parents removed; 3.Without parental care
(brood mass not present and parents removed).
13.14 +/- 0.45
11.40 +/-0.30
10.20 +/- 0.37
P<0.05
Contd..
The larval development required is 10.20 +/- 0.37 ,11.40 +/-0.30 &13.14 +/- 0.45
days respectively in Group 1,2 & 3
Survival of larva at L1 is 97%,95% &90% respectively
Treatment groups
Offspring
survival
Parameter
Ni
L1
L2
L3
1) Control: parental care
390
97
90
70
2) With Brood-mass. Parents removed prehatching
468
95
89
60
3) Without parental care : with fresh meat,
brood-mass no present and parents
removed pre-hatching
351
90
77
28
Contd.
Significant differences in the variation the number of offspring surviving to adult
emergence between three experimental groups
The number of adults that emerged was significantly different among the three
groups (X2 =11.96, df = 5, P , 0.05)
Compared
groups
Difference
1 Vs 3
28.13 12.78 = 15.35
3.214
1 Vs 2
28.13-25.93 = 2.2
0.47
2 Vs 3
25.93 -12.78 = 13.15
2.75
Contd.
Contd.
Larval development was significantly shorter in the control group
The presence of both parents and brood mass were required for larvae to reach adulthood
The differences is due to better nutritional quality of the food prepared by parents through
the oral and anal secretions
Case study 2
Maternal vibration induces synchronous hatching
in a sub- social burrower bug
By-Hiromi et al., 2012
Objective :-
To investigate the effect of mother presence on hatching synchrony
What effect does parental care have upon hatching of embryos?
To ascertain whether embryos hatch , in response to the vibrational signal of mother
Result & Discussion
Occurrence pattern of maternal vibration. Means +SD are shown for bouts/min from the start of vibration
generated by seven mothers.
Hatching was synchronous, passing through a fixed series of sequential steps
Hatching pattern of embryos exposed to the different treatments. Mean + SD are shown for the percentage of
hatched eggs at 5 min intervals for the first 90 min, and for 12 h and 24 h from the start of hatching.
47.8+/-17.4 bouts of vibration /min
13.7+/-1.4 bouts /min
Schematic diagram showing artificial vibration. Coiled a rigid wire around an electrodynamics cordless motor
glued a dead mother touching a mature egg mass with mid-leg
92.0+/- 13.7
88.2 +/- 14.4
b
39.4+/- 22.2
80.8 +/- 17.6
Duration of hatching
Rate of vibration
At 1st minute
13.7+/-14.4 bouts of vibration /min
At 8th minute
47.8+/-17.4 bouts of vibration /min
At the end
39.1+/-13.8 bouts of vibration /min, Mother Starts trophic egg production
Duration of hatching
Hatching success in Control & treated group
After 12 hour
97.0 +/- 4.5% and 94.5 +/- 6.5% respectively
After 24 hour
97.0 +/- 4.5% and 96.0 +/- 3.5% respectively
Duration of hatching
Hatching success during artificial vibration
At 1st 15 minute
39.4+/- 22.2
After 90 minute
80.8 +/- 17.6
After 12 hour
88.2 +/- 14.4
After 24 hour
92.0+/- 13.7
Parents regulate the hatching.
Mothers were observed to guarding the egg mass without feeding for 6 to7 days
Highly synchronous hatching begins at 8.7 +/- 1.4 , which corresponded to the peak of the
maternal vibration
CONCLUSION
Parental care have many adaptive advantage
for offspring
Synchronous hatching promote aggregation of
young-one
Large scale aggregation of offspring act as
cooperative defense
FUTURE PROSPECTS
Study of parental care behavior help in mass killing of insect by
locating the brood site
If it is a parasitoid of any insect pest ,we can colonies it by habitat
management to control the host species