Development

Insect and Development Study Guide

Key Points:
- Understand the difference between metamorphosis types.
- Know the characteristics of various larval forms
- Recognize the importance of imaginal discs in holometabolous development
- Familiarize yourself with the embryonic development stages

I. Early development:
A. Egg structure:
1. Chorion: the outer shell of the egg
2. Micropyles: passages for sperm entry
3. Aeropyles: passages for gas exchange
4. Yolk: Contains lipids, proteins, and carbohydrates
B. Fertilization process:
1. Stored sperm released onto eggs as they pass down the oviduct
2. Multiple sperm may enter the egg (polyspermy)
3. Only one sperm fertilizes the egg
C. Blastoderm formation
1. Nuclear division without cytokinesis (membrane formation), producing a
syncytial blastoderm - Nuclei are totipotent (cells are capable of giving
rise to any cell type)
2. Cell membrane formation around each nucleus and migration to the
periphery to form a single layer
3. Germ bad formation- the remainder of blastoderm cells form a thin
membrane (serosa) around the yolk
4. The membranous portion of blastoderm folds over and encloses the germ
band
5. Germ band differentiates into ectoderm and mesoderm
D. Embryotic development:
1. Segmentation becomes apparent
2. Endoderm forms the gut (before hatching, the foregut and the hindgut
meet to form the alimentary canal)
3. Ectoderm differentiates into integument, tracheal system, sense organs,
and nervous system
II. Postembryonic development:
1. Sexually immature stages called larvae
2. Stages between molts called instars
3. The adult stage is referred to as the imago (growth stops at maturity
except in apterygotes and Ephemeroptera)
4. Functional wings (if present) only in adult stages ( some Ephemeroptera,
sexually immature winged instar, the subimago, molts into winged adult)
5. Imaginal discs: undifferentiated cell groups in holometabolous larvae,
grow into adult tissues during pupation
III. Patterns of Metamorphosis
 A. Ametabolous
1. Continuous molting after maturity
2. Offspring and adults are ecologically similar
B. Hemimetabolous (incomplete “punctuated” metamorphosis-aquatic)
1. Aquatic juveniles called naiads
2. Juveniles are ecologically different from adults
a) Odonate naiads are predatory and have modified labium that
extends to capture prey
3. Examples: Odonata, Ephemeroptera, Plecoptera
C. Paurometanoulous: (incomplete metamorphosis ”gradual” -terrestrial)
1. Terrestrial juveniles called nymphs
2. Juveniles are ecologically similar to adults
3. Examples: Hemiptera, Orthoptera
D. Holometabolous: (Complete metamorphosis)
1. Various larval forms
2. Juveniles are ecologically different from adults
3. Includes pupal stage
4. Examples: Lepidoptera (caterpillar), Diptera (maggots), Coleoptera
(grubs)
IV. Types of endopterygote larvae
A. Vermiform
1. No legs, poorly sclerotized
2. Found in soil, dung, decaying tissues
3. Vulnerable to desiccation and predation
4. Examples: Hymenoptera, Coleoptera, and Diptera
B. Campodeiform:
1. Functional thoracic legs, prognathous mouthparts
2. predatory, sclerotized
3. Examples: Neuroptera, Coleoptera, Trichoptera
C. Scarabaeiform
1. Grub-like, C shaped
2. Short, sclerotized thoracic legs
3. Examples: some Coleoptera (scarab beetles)
D. Elateriform
1. Worm-shaped with heavily sclerotized bodies
2. Short thoracic legs
3. Example: some Coleoptera (wireworms)
E. Eruciform
1. Caterpillar-like
2. Thoracic legs and abdominal prolegs
3. Examples: Lepidoptera (up to 5 pairs w/ crochets “hooks”), some
Hymenoptera (6+ pairs, no crochets, sawflies)
Questions:
Are these descriptions of larval forms of any use in phylogenetic reconstructions?
- No, there are too many convergent evolutionary designs. It may help with identification
and ecological needs.
Diapause & Thermorgulation
Study Guide: Diapause & Thermoregulation
Key Concepts:
- Comparison between quiescence and diapause
- Mechanisms of cold tolerance in insects
- Adaptations for heat and aridity survival
- Importance of environmental cues in insect life cycles.

1. Generation times and seasonability:

a. Voltinism: Number of generations per year
i. Univoltine: one generation per year
ii. Bivoltine: two generations per year
iii. Multivoltine: multiple generations per year
iv. Long maturation and adult lifespans
2. Seasonality and Arrested Development (summer/winter & dry/wet) :
a. Quiescence
i. Arrested or slowed development in direct response to unfavorable
conditions
ii. Development resumes as soon as conditions improve
b. Diapause
i. Prolonged arrested development regulated by hormones
ii. Corresponds to seasonal changes
iii. May occur at any stage of development
● Obligatory diapause:
a. Occurs at a fixed time in development
b. Common in univoltine insects
i. The life cycle of a Mountain Pine Beetle;
1. August-> adult lays eggs -> larvae until
November-> Diapause starts as 4th instars
-> may a pupa forms-> adult-> fuck then
start over
● Facultative diapause:
a. Occurs only in generations that must survive unfavorable
conditions
b. Found in bi- or multivoltine species
3. Diapause induction and breaking
a. Environmental cues: day length, temperature, food, etc.
b. Costs of early diapause entry:
i. Size limitation, no food in winter, too cold
c. Benefits of early diapause entry:
i. When fall comes around, the insect will be bigger and have more time to
find food & reproduce
d. Importance of multiple cues for diapause decisions
i. Single cues may be less informative of risk
 ii. Day length is correlated w/ temperature but not perfectly. Natural
selection favors the correct decision
e. Selective pressures on diapause timing
4. Thermoregulation
a. Ectothermy:
i. Heat derived from the external environment
ii. Basking to elevate internal temperature
b. Endothermy
i. Heat gained from energetic metabolism
● Access to high-energy foods may engage in limited endothermy
● Basking combined w/ rapid shivering of flight muscles generates
heat needed for flight
● The thoracic temperature may exceed ambient by 20-30° C
ii. Examples: insects with insulating hairs/scales, honey bees-remain
clustered in the colony and derive heat from the metabolism of stored
honey
5. Dealing with Cold
a. Freeze-susceptible insects: these insects will die if their tissues freeze
i. Migration
ii. Endothermy
iii. Chill tolerance: Water and snow can provide insulation from cold.
b. Freeze avoidance (using glycerol to lower freezing point)
i. If the temperature drops below a new freezing point, the insect dies
c. Freeze-tolerant insects
i. Can survive periods of freezing
ii. Seasonal production of ice nucleating agents
iii. Partial cell dehydration to reduce intracellular freezing risk
● Freezing is encouraged outside of cells by proteins and crystalline
substances in hemolymph and gut
6. Dealing with heat and aridity
a. Heat adaptations:
i. Protein denaturation at high temperatures
ii. Acclimation (gradual exposure to changing temperatures) to reduce
mortality relative to instantaneous exposure
iii. Behavioral mechanisms (e.g., Burrowing, nocturnal activity)
iv. Morphological adaptations to reduce heat gain from hot surfaces
b. Aridity adaptations:
i. Physiological adaptations to reduce water loss
ii. Behavioral adaptations (e.g., fog basking)
Nutrition & Digestive system
Study guide: Digestion and Nutrition
Key Points:
- Digestion typically occurs in the midgut
- Absorption mainly takes place in the midgut
- Nitrogenous waste is excreted as uric acid (water-conserving but energetically
expensive)
- Malpighian tubules and rectal pads work together for waste elimination
I. Nutritional Ecology
A. Symbiotic microorganisms
1. Common in insect guts, especially when diet lacks essential nutrients or
contains indigestible components
2. Examples:
a) Leaf-cutter Ant
(1) Foragers (mediae) drag vegetation back to their nests;
Minims tend to fungus gardens and macerate leafy tissue
to inoculate it with enzymes in feces
(2) Cultivate mutualistic fungus in underground nests; fungus
no longer produces spores (fully domesticated)
(3) Fungus produces gongylida for larval feeding
b) Blood feeders
(1) Obligate (e.g., bed bugs) and facultative (e.g., mosquitoes,
lice)
(a) Facultative blood-feeder females require a blood
meal for egg production; some essential nutrients
are stored from larval stages; adults may also feed
on nectar
(2) Blood is rich in proteins, lipids, irons, and specific vitamins.
Internal symbionts provide essential nutrients lacking in
blood (vitamin B and other nutrients)
c) Termites
(1) Rely on gut symbionts (protozoans/bacteria) to digest
cellulose and release sugars; animals cannot produce
cellulase to break down woody material:
B. Transmission of internal symbionts
1. Oral uptake via anal and oral secretions
2. Intracellular microorganisms in mycetocytes or bacteriocytes: Symbiots
transferred to the eggs or embryos before parturition
II. Essential Nutrients
A. Water:
1. Uptake mechanisms: drinking, eating, oxidative metabolism (the chemical
process that uses oxygen to make energy from carbohydrates, water
vapor
B. Carbohydrates:
 1. Important for flight energy and chitin synthesis (chitin is a polysaccharide,
which is a chain of sugars)
C. Amino Acids (Nitrogen):
1. Higher in animal prey, lower in vegetation
D. Lipids:
1. Energy storage as fat
2. Sterols required for hormones (e.g., ecdysone)
E. Vitamins:
1. Must be derived from diet
F. Minerals
1. Insects require less calcium and iron than vertebrates; there is no
skeleton, so there is less need for calcium, and there is no hemoglobin,
so there is not as much need for Iron.
III. Digestive system
A. Foregut: (storage, pre-digestion, grinding)
1. Mouth: food manipulation, saliva production
2. Esophagus: extends through the thorax
3. Crop: food storage
4. Proventriculus: controls food release, grinds food)
B. Midgut: (digestion and absorption) (Hymenoptera: the midgut is a blind sac until
pupation)
1. Ventriculus: a tube-like sac that secretes all necessary digestive enzymes
2. Gastric cecae: fingerlike projections that house symbiotic bacteria needed
for digestion. Larger in herbivores are because plants are more
challenging to digest and need more time and surface area to digest
nutrients fully.
C. Hindgut (reabsorption and Waste removal)
1. Malphighian tubules: remove nitrogenous waste (~Kidney)
a) Water, salts, sugars, amino acids, and nitrogenous waste pass
into the malphighian tubules via osmosis and active transport.
b) Ammonia is converted to urea and then uric acid, energeticall
expensive but conserves water.
2. Ileum & rectum: selective reabsorption of water, amino acids, sugars, and
ions. Most reabsorption of water (necessary for homeostasis) occurs in
rectal pads
3. Anus: passes fecal pellets
Respiration and Circulation
Insect Respiration System:
I. Tracheal system:
A. The primary system for gas exchange in insects
B. Components:
1. Spiracles: external openings (up to 10 pairs)
2. Tracheae: Tubes extending from spiracles
3. Tracheoles: Primary sites for gas exchange to cells
4. Air Sacs: enlarged dilations of tracheae used in ventilation
II. Spiracles:
A. located on thoracic (2-3) and abdominal segments (up to 8)
B. Musculated and capable of closing
C. Airflow pattern: in through anterior, out through posterior spiracles
III. Tracheae:
A. Tubular network, historically similar to integument
B. Reorganized during metamorphosis to accommodate demands for oxygen by
flight muscles but basic design remains unchanged.
IV. Ventilation
A. Improves air circulation in the tracheal system
B. Methods:
1. Abdominal pumping in grasshoppers
2. Thoracic movements during flight and open anterior spiracles provide
flight muscles with O2
C. Increases water loss due to air saturation in the tracheal system
V. Gas exchange in aquatic insects
A. Methods:
1. Diffusion across the body (small insects)
2. Closed tracheal system (gills in aquatic nymphs)
a) Gills (modified trachea) often employed
3. Open tracheal systems:
a) Direct connections with the atmosphere (e.g., mosquito larvae)
(1) Only terminal spiracles open
b) Connections to air pockets in aquatic plants
4. Air bubbles (physical gills)
a) Physical gills: Passive diffusion replenishes the oxygen supply as
the animal breathes. The insect can remain underwater as long as
the O2 concentration is greater than the O2 that it uses.
b) The surface area of the bubble is positively correlated with
diffusion efficiency. Insects cannot stay underwater indefinitely due
to nitrogen diffusing out of the bubble.
c) Trapping water among setae and constraining bubble size;
nitrogen diffusion out of the bubble, partial pressure changes
cause O2 to replace the lost N
5. Hemoglobin (rare, found in only three cases)
a) Bloodworms: live in oxygen-deficient water
 b) Horse bot fly: Larvae live in the stomach of host; Hemoglobin
allows uptake of O2 when a gas bubble passes by
c) Backswimmer: uses hemoglobin in the abdomen to extend and
stay underwater
Insect Circulation system
I. Open circulatory system
A. Hemolymph percolates through the hemocoel (body cavity)
II. Components:
A. Dorsal Blood Vessel (DBV)
1. Heart: an abdominal region with ostia (valve-like perforations)
2. Aorta: section w/o ostia
B. Characteristics:
1. Not associated with O2 or CO2 transport
2. Almost no blood pressure
3. Complete blood mixing takes 5-30 minutes (vs. 2-4 minutes in humans)
C. Functions:
1. Transportation of:
a) Nutrients
b) Waste products
c) Hormones
2. Provides hydrostatic pressure for molting and locomotion
3. Immune response
4. Thermoregulation
Key differences from Vertebrates:
1. The insect circulatory system is open, not closed
2. Not primarily associated with respiration
3. Uses hemolymph instead of blood
Locomotion and Flight
Insect muscles, locomotion, and flight
1. Insect muscles:
a. Characteristics:
i. All insect muscles are striated
ii. Located interior to the skeleton
iii. Attached to exoskeleton via tonofibrillae (secretions of epidermal cells)
rather than tendons
iv. Apodemes often serve as muscle attachment sites
v. Strength:
1. Similar to vertebrate muscles
2. “Feats of strength” result of small size and effective use of levers
3. Allometric scaling governed by square-cubed law
2. Insect Joints:
a. Monocondylic (ball and socket)
i. Greater range of motion
ii. Less stability
b. Dicondylic (hing)
i. Limited range of motion
ii. Greater stability and strength
c. Examples:
i. Body-coxa and coxa-trochanter: alternating dicondylic joints
ii. Femure-tibia: Dicondylic
iii. Tibia-tarsus: often monocondylic
3. Insect wings
a. Variations among insects:
i. Dragonflies: independent wing control
ii. Lepidoptera and Hymenoptera: joined front and back wings
iii. Some insects: modified wings (e.g., flies halteres for balance)
iv. Beetles: forewings as protective covers
4. Flight mechanisms:
a. Direct flight mechanisms (Paleoptera)
i. Wings operate independently
ii. Dorsalventral muscles act directly on wings
b. Indirect flight mechanisms (Neoptera)
i. Muscles connect to the thorax, not wings
ii. Upstroke and downstroke are controlled by thorax deformations
iii. Longitudinal muscles contract, causing the notum to rise for downstroke
iv. Dorsalventral muscles contract, causing notum to lower for upstroke
5. Wing movement control:
a. Direct flight muscles are still involved in indirect flight mechanism, muscles
control pronation and supination
b. Basalare contractions: lowers leading edge (pronation)
c. Sublare contraction: lowers trailing edge (supination)
6. Evolution of wings:
 a. Evolved once in insects
b. Veins and articulatory sclerites homologous among pterygotes
c. Six-legged condition evolved before wings
7. Origin of wing theories: (wings for powered flight have evolved 4 times in the hx of life-
Pterosaurs, birds, bats and insects: unlike vertebrates, insects did not have to sacrifice a
pair of limbs for wings)
a. Paranotal theory:
i. Wings evolved from lateral expansions of terga, provided stability for
gliding; development and refinement of basal hinges and musculature
would complete the process
ii. Supported by broad thoracic nota in fossil Ephemeroptera
iii. Criticism: articulating structures would need to evolve de novo (from
nothing)
b. Exite theory:
i. Wings evolved from mobile thoracic pleural structures, either gill plates or
coxopodites (basal leg structures)
ii. Supported by fossil nymphs gill-like structures: moveable, thin
membranous, similar venation to wings, basal articulation present
iii. Molecular evidence links exite expression in crustaceans with insect
wings
1. Wings and external gills share common underlying developmental
pattern of gene expression
8. Origin of Flight hypothesis:
a. Aquatic origin:
i. Surface-skimming theory: rudimentary wings for thrust on water surfaces
ii. Bridges gap between gill flapping and wing flapping
b. Terrestrial origin:
i. Gliding theory: flight arose in arboreal insects
ii. Recent phylogenetic analysis suggests terrestrial common ancestor for
pterygotes
iii. Gliding behavior observed in primitive wingless insects (Archeaognatha)
Key points to remember:
- Insect muscles are striated and attached via tonofibrillae
- Understand the differences between monocondylic and dicondylic joints
- Know the direct and indirect flight mechanisms
- Familiarize yourself with the theories on wing origin and flight evolution