Title: Learning and Memory across the Lifespan
1Chapter 12
- Learning and Memory across the Lifespan
212.1 Behavioral Processes
312.1 Behavioral Processes
- The Developing Memory Infancy through
Adolescence - Learning and Memory in Everyday Life Can
Exposure to Classical Music Make Babies Smarter? - Sensitive Periods for Learning
- The Aging Memory Adulthood through Old Age
4Developing Memory in Infancy Some Learning Can
Occur Before Birth!
- Gestational age (GA)time since conception.
- By 25 weeks GA, enough development in fetuss
brain and sense organs to perceive. - In studies (play sounds to human fetus)
- Fetus (3436 weeks GA) moved in response to a
sound habituated (reduced response) by trial 13. - Moved to 2nd stimulus habituated by trial 11.
- Back to 1st stimulus habituated by trial 8.
5Habituation to Sound (in 10 Human Fetuses)
Adapted from Hepper Shahidullah, 1992.
6Time (sec.)
Polygraph recording of sucking on nipple
Figure adapted from Figure 1 of DeCasper
Spence, 1986.
Short pause
Long pause
Long pause
Unfamiliar story plays
Familiar story plays
Familiar story plays
Details of DeCasper Spence (1986)
paradigm Before birth, infants were played
familiar story in mothers voice memory for
story was tested after birth by playing
recordings of this story or unfamiliar story
while infants sucked on artificial nipple.
Infants tend to suck in bursts punctuated by
pauses (interburst intervals, IBIs). First,
researchers took baseline measurements of average
IBI. Then, conditioned some infants that long
IBIs would be reinforced with familiar story
while sucking short IBIs would be punished with
unfamiliar story while sucking. (Other infants
in counterbalanced conditioning, short intervals
were punished.)
7Conditioning and Skill Learning in Young Children
- Explosion of learning in first few years of life!
- Most learning present in adults, present in
infants. - But, perceptual and motor systems immature.
- Until input/output systems mature, infants
cannot fully learn or express.
8Rovee-Collier Studies
- (1993) Rovee-Collier studied instrumental
conditioning in infants - 2-month-old infants learned to kick to move a
colorful mobile (hung over the crib). - Illustrates instrumental conditioning.
- With no reminders, Infants remembered foot-kick
technique for 13 days. - With reminders, up to 21 weeks.
9Rovee-Collier Studies
- If crib liner with new pattern was used, babies
didnt kick. - Illustrates context-dependent learning.
Courtesy of Carolyn Rovee-Collier
10Infants and Classical Conditioning
- Other studies show infants have basic components
of classical conditioning. - Human and rat infants learned delay eyeblink
conditioning. - But, use more trials than adults of their
species. - Trace conditioning improved from infancy to
early adulthood. - Shows that, with more mature development,
organism can learn more efficiently (under
increasingly difficult conditions).
11Development of Episodic and Semantic Memory
- Elicited imitationinfants ability to imitate an
action at a later time. - From single observational learning training
session. - In study, 10-month-olds are shown how to operate
a toy puppet. - 4 months later, showed more interest in the
puppet than control group (same age, no prior
showing). - At 5 years, showed more interest and dexterity
with the puppet than control group, though most
could not recall previous exposure.
12Development of Episodic and Semantic Memory
- In study, 4-, 6-, and 8-year-olds taught 5 facts
from an experimenter and 5 from a puppet. - One week later, 6- and 8-year-olds recalled and
recognized more facts than 4-year-olds. - 6- and 8-year-olds had better recall of source,
with more intra-experimental errors (i.e., knew
it was learned in experiment, confused source). - 4-year-olds made more extra-experimental errors
(i.e., thought learning was outside experiment,
for example at school).
13Episodic Memory in Children
Data from Drummey Newcombe, 2002.
14Development of Working Memory
- Working memory lifespan progression
- English-speaking children 56 years can hold
average digit span of 34 digits in working
memory. - By 910 years, can hold 56 digits.
- By 1415 years, can hold 7 digits (adult
average). - Similar working memory progression seen with
words and visual patterns. - Why fewer for children?
- Lack of exposure.
- Childrens performance improves with familiarity.
159
8
7
6
Girls
5
Mean Digits
Boys
4
3
2
1
0
5
6
7
8
9
10
11
12
13
14
15
Age (years)
Memory for digit span increases with age (reach
adult levels by about age 12 or 13) no
significant gender difference. Figure plotted
from data in Gardner, R. (1981). Digits forward
and digits backward as two separate tests
Normative data on 1567 school children. Journal
of Clinical Child Psychology, Summer 1981,
131135.
16Learning and Memory in Everyday Life Can
Exposure to Classical Music Make Babies Smarter?
- Limited intellectual benefits from exposure to
classical music (no true Mozart effect). - Research shows little evidence that supports
benefits zero evidence that the effect lasts
longer than 1015 minutes. - So, why did scores increase?
- Music may prime or prepare brain regions for
abstract spatial reasoning or mental imagery. - Music may improve mood and subsequent
performance.
17Sensitive Periods for Learning
- Sensitive periodstime ranges during which
learning is enhanced or possible. - Examples
- In male sparrows, 30100 days is a sensitive
period for song learning. - In cats, 3 weeks to 60 days is a sensitive
period for visual development. - But, for monkeys, all of the first 6 months are
important for visual development.
18Sensitive Periods for Learning
- In study of 28 human infants who had cataract
surgery at age 1 week to 9 months - ACUITY improved significantly over 1 month,
with some improvement apparent after as little as
1 hour of visual input. - Unlike older children, improvement was the same
for eyes treated for monocular and binocular
deprivation. - Visual input necessary for postnatal
improvement its onset initiates rapid functional
development.
19Imprinting
- Imprintingphenomenon in which some species
(e.g., newborn goslings, turkeys, sheep, deer,
buffalo) form a social bond with the first
object they see. - Imprinting involves critical period for
permanent change to occur. - http//www.youtube.com/watch?vLGBqQyZid04
Thomas D. McAvoy/ Time Magazine
20Social Attachment Learning
- Primates do not appear to imprint, but there is
evidence of sensitive period for social
attachment. - In study, Harry Harlow rears rhesus monkeys
isolated from mothers in adolescence moved to
group cages, show social retardation. - For rhesus monkeys, first months sensitive
period for learning social interactions. - http//www.youtube.com/watch?vAn02zCsVEpY
21Social Attachment Learning
- Sensitive period for social attachment in humans?
Consider children under Ceausescu regime (1970s) - Romanian children (RC) reared from infancy (up
to 42 months) in depriving institutions, then
placed in UK adoptive homes. - Compared with nondeprived UK-born children
adopted before 6 months. - past
- http//www.youtube.com/watch?vrXivHuugp3c
- present
- http//www.youtube.com/watch?vFWKQNMZa--Yfeature
related
22Social Attachment Learning
- Findings
- RC tested at time of entry to UK showed
developmental impairment in cognitive function. - Tested again at 4 years all RC show
improvement. - RC adopted before aged 6 months showed normal
cognitive and social functioning. - But, RC adopted at 6 months or later still showed
some cognitive deficits, mild social problems. - Suggests biological programming or neural
damage from institutional deprivation varied
outcomes related to early environmental
stimulation.
23A Sensitive Period in Humans
Data from Rutter et al., 1998.
24Aging Memory Adulthood to Old Age
- Working memory capacity is particularly
vulnerable in old age. - Average STM capacity for digits drops from 7 (in
early to middle adulthood) to 66.5 in elderly
adults.
25Conditioning and Skill Learning DeclineBut
Well-Learned Skills Survive
- Learning decline begins around age 4050 in
humans. - Also seen in elderly rabbits, rats, and cats.
- However, well-established, highly practiced
skills tend to be maintained or improved (chess
and bridge experts).
26Conditioning and Skill Learning DeclineBut
Well-Learned Skills Survive
- In studies
- Eyeblink conditioning may take twice as many
trials in elderly. - Skill learning for rotary pursuit task and
computer use take more time.
27(B) Rotary pursuit skill learning
(A) Eyeblink classical conditioning
60
50
Mean distance covered
40
Trials to Criterion
30
20
10
0
18-29
30-39
40-49
50-59
60-69
70-70
Age (in years)
Age (in years)
Classical conditioning and skill learning decline
with aging.
(A) Plotted from data in Solomon et al., 1989,
Table 1. (B) From Kausler, D. (1994). Learning
and Memory in Normal Aging. New York Academic
Press, p. 38 fig 2.4 (top), which cites adapted
from Ruch, 1934.
28Episodic and Semantic Memory Old Memories Fare
Better than New Learning
- Healthy elderly adults tend to retain semantic
knowledge, and recall many episodic memories. - In paired associates test
- Elderly may be able to recognize words or
images previously studied. - May have difficulty with recall.
- May recall more information if given more time
or allowed to self-pace rate of presentation.
29Paired associate learning is impaired in elderly
adults relative to young adults when items are
presented at a rate of one every 1.5 seconds
impairment decreases if presentation rate is
slowed. Recall best when learning is self-paced,
though elderly subjects never quite reach same
performance as young subjects. From D. Kausler
(1994) Learning and Memory in Normal Aging, NY
Academic Press, p. 88, which cites adapted from
Canestrari, 1963, Table 2.
3012.1 Interim Summary
- Just about every kind of learning and memory
observed in adults can also be observed in very
young children. - Some simple kinds of learning (e.g.,
habituation, recognition) can be observed before
birth. - Other kinds of memory (particularly episodic
and working memory) may be present at a very
young age, but do not fully mature until late
childhood or adolescence.
3112.1 Interim Summary
- Development of learning and memory abilities at
least partially reflects brain development. - Sensitive periods time windows early in life
when certain kinds of learning advance most
rapidly. - Includes imprinting, social attachment learning.
3212.1 Interim Summary
- Many kinds of learning and memory show some
decline in healthy aging. - Working memory is especially vulnerable.
- In other memory domains (e.g., skills,
conditioning, episodic and semantic memory) old,
well-formed memories tend to survive well may be
harder to acquire new memories.
3312.2 Brain Substrates
3412.2 Brain Substrates
- The Genetic Basis of Learning and Memory
- Neurons and Synapses in the Developing Brain
- Gender Differences in Brain and Behavior
- The Brain from Adulthood to Old Age
35The Genetic Basis of Learning and Memory
- DNAmaterial in cell nucleus instructions for
replication. - Looks like twisted ladder sides sugar and
phosphate molecules, rungs base pair. - Four kinds of DNA
- Adenine
- Thymine
- Cytosine
- Guanine
36The Genetic Basis of Learning and Memory
- DNA organized into chromosomes.
- Humans have 23 chromosome pairs (one set from
each parent). - 23rd pair determines gender.
- XX female
- XY male
- Chromosomes subdivided into genes segment of DNA
with information for building proteins from amino
acids. - Probably 20,000 to 25,000 genes in humans.
37Genes and DNA
CNRI/Photo Researchers, Inc.
38Genetic Variation among Individuals Affects
Innate Learning Abilities
- Mutationaccidental changes in DNA sequence.
- Possibly from outside causes (e.g., radiation,
viral infection) or copying error. - Mutations can
- Be harmless.
- Lead to cell malfunction, disease, death.
- Be beneficial to the species.
- New characteristics for reproduction or survival.
39Genetic Variation among Individuals Affects
Innate Learning Abilities
- Because of mutation over time, most genes have
allelesnaturally occurring variations. - e.g., eye color
40Genetic Variation among Individuals Affects
Innate Learning Abilities
- Brain function also influenced by variations can
affect learning and memory. - Examples
- BDNF protein (the Val allele) may facilitate
long-term plasticity. - Tyr allele (variant of His allele on 5-HT2AR
gene) results in less-efficient serotonin
receptors. - Perform slightly worse on delayed word recall
task.
41Genetic Influences on Learning and Memory in
Humans
(a) Data from Egan et al., 2003 (b) adapted from
de Quervain et al., 2003.
42Selective Breeding and Twin Studies
- Tryon (1940) Can animals be bred for learning
ability? - Bred discrete groups of maze-bright and
maze-dull rats in 7 generations. - By 7th generation, maze-bright offspring
routinely out-perform rats bred from maze-dull
line.
43Data shown are hypothetical, based on Tryon, 1940.
44Selective Breeding and Twin Studies
- Multiple genes control characteristics of
learning ability. - No single gene.
- Human twin studies suggest that over half of the
variation in memory scores may be genetic. - Identical twins have more similarity than
fraternal.
45The Influence of Environment
- Rats raised in enriched environment (good
sensory stimulation) have more dendrites and
synapses. - Males had the most growth in visual cortex.
- Female rats had the most growth in frontal
cortex.
46Neurons and Synapses in the Developing Brain
- Neurons are overproduced, then weeded out.
- Neurogenesis (neuron birth) most active during
prenatal development continues to a limited
degree throughout life. - Not uniform throughout brain some neurons form
earlier than others.
47Neurons and Synapses in the Developing Brain
- In early development, glia guide cell migration
produce molecules that modify growth of axons and
dendrites. - Some glia (oligodendrocytes) produce myelin
sheath, from birth to 18 years. - Neurotrophic factors (e.g., BDNF protein) help
cells properly locate and specialize. - Without these chemical compounds, about 1/3 of
neurons die (apoptosis), a natural phenomenon.
48Neurons and Synapses in the Developing Brain
- Synapses are also formed, then pruned.
- Synaptogenesis (formation of new synapses)begins
during gestation, but most active after birth to
about age 6. - Tiny dendrite spines come and go if stimulated
by neurotransmitters, synapses may form. - Unused synapses die (pruning).
- New synapses may strengthen during non-REM
sleep and unused may die during REM sleep.
49Most synapses Occur on Dendritic Spines
(a) Adapted from Hof Morrison, 2004 (b)
adapted from Trachtenberg et al., 2002.
50Sensitive Periods for Learning Reflect Sensitive
Periods for Neuronal Wiring
- Neural pathways (and specific receptors) may
develop rapidly during sensitive periods. - Apoptosis may then clean up neurons not used in
in this sophisticated development.
51The Promise of Stem Cells for Brain Repair
- Young brains highly plastic older brains less
able to adjust. - Can stem cells be integrated into adult brains?
- Stem (especially from fetal tissue) cells have
ability to develop into many cell types. - e.g, skin, liver, brain cells
- Fetal stem cell transplant research still
preliminary. - Tried in Parkinsons disease patients.
- New neurons do not cure the underlying disease.
52Embryonic Stem Cell Transplants in Brains of
Parkinsons Patients
Adapted from Freed et al., 2001.
53Gender Differences in Brain and Behavior
- In studies
- Women often perform better than same-aged men
on - List recall.
- Story recall.
- Memory for object location.
- Men can outperform women in maze learning.
- Men and women studied a fictitious town map
- Men tended to learn a route more easily.
- Women remembered more landmarks.
54Gender Differences in Brain and Behavior
- Male and female rats also show gender
differences. - Sex hormones may contribute to gender-based
learning differences.
55Effects of Sex Hormones on Brain Organization
- Pubertybodys physical change to sexual maturity
in adolescence. - Surge in release of sex hormones.
- Primarily estrogens in woman, androgens in men
(especially testosterone). - In mammals and birds, testosterone surges in
female fetuses and even more in male fetuses just
before birth.
56Effects of Sex Hormones on Brain Organization
- During infancy, testosterone influences sex
differences in brain development. - Larger in women
- Lateral frontal cortex
- Language areas (supramarginal gyrus)
- Hippocampus
- Larger in men
- Visual and spatial processing areas
57Effects of Sex Hormones on Adult Behavior
- Gender differences in memory performance appear
after puberty from circulating estrogen and
testosterone. - Estrogen stimulates adult rats neuronal growth
and synaptic plasticity (LTP), especially in the
hippocampus.
58Effects of Sex Hormones on Adult Behavior
- Estrogen may increase verbal learning
testosterone may increase spatial learning. - But, relationship between sex hormones
(especially testosterone) and learning is
complex. - Studies show male-to-female transsexual persons
taking estrogen scored higher on paired-associate
task. - Compared to similar group who had not yet
started estrogen treatment.
59Adulthood to Old Age Parts of the Aging Brain
Lose Neurons and Synapses
- Slow human brain shrinkage, including the
cerebellum, begins in young adulthood. - By age 80, average adult loses about 5 percent
of brain weight. - Studies show
- Cerebellum-dependent classical eyeblink
conditioning slows with age. - However, there is little loss of hippocampal
neurons in the healthy elderly. - Reductions in neurons disease warning signs.
60Neuron Loss in Prefrontal Cortex of Aging Monkeys
Adapted from Smith et al., 2004.
61Synaptic Connections May Be Less Stable in Old Age
- Barnes (et. al.) suggest total number of neurons,
synapses does NOT decrease rather, decrease in
ability to maintain changes in synapse strength. - Rat and monkey studies suggest that synapses
may be less stable in old age. - In studies, young rat and an old rat learned a
figure 8-shaped maze. In second session,
hippocampal LTP in the old rat was unstable. - Instability could contribute to spatial and
episodic memory declines.
62Hippocampal Neurons Encoding Location in Old and
Young Rats
(be) adapted from Barnes et al., 1997.
63New Neurons for Old Brains? Adult Neurogenesis
- Adult brain may be able to grow new neurons.
- Adult neurogenesis has been studied (and
reliably observed) in birds, fish, amphibians,
reptiles. - Neurogenesis in mammals?
- Studies show limited neurogenesis in brains of
adults macaque monkeys and human cancer patients. - Most new neurons die within a few weeks.
6412.2 Interim Summary
- Development of learning and memory abilities at
least partially reflects brain development. - Temporal and frontal cortex are among the last
brain areas to fully mature. - May help explain why memory processes dependent
on these areas are among last to reach full adult
potency.
6512.2 Interim Summary
- Genes play a large role in determining learning
and memory abilities. - Enriched environment studies show that
experiences can also impact brain organization
and an individuals abilities. - Before birth, the brain overproduces neurons and
synapses. - Unnecessary neurons and synapses are gradually
eliminated.
6612.2 Interim Summary
- Sensitive periods may reflect times when external
inputs can easily and profoundly alter brain
connectivity. - After sensitive period, large-scale
organization of brain area in question may be
fixed, and further learning (of the kind in
question) may be limited to fine-tuning.
6712.2 Interim Summary
- Sex hormones, like estrogen and testosterone, can
influence development and performance. - Leads to gender differences among adults in
various kinds of learning and memory. - Influence on developing brain leads to gender
differences even in very young individuals.
6812.2 Interim Summary
- Working memory declines in healthy aging.
- Vulnerability may reflect normal frontal cortex
shrinkage in healthy aging. - Pattern of memory loss in healthy aging may
reflect loss of neurons and synapses. - Also, may reflect decrease in ability to
maintain changes in synapse strength. - Thus, newly encoded information may be lost.
6912.2 Interim Summary
- New neurons produced throughout the lifespan.
- But, particularly in humans, there is as yet
little evidence that adult neurogenesis could
provide large-scale replacement for damaged or
aging neurons.
7012.3 Clinical Perspectives
7112.3 Clinical Perspectives
- Down Syndrome
- Alzheimers Disease
- A Connection between Down Syndrome and
Alzheimers Disease? - Unsolved MysteriesTreating (and Preventing)
Alzheimers Disease
72Down Syndrome
- Down syndromecongenital form of mental
retardation which occurs equally in girls and
boys. - Retarded speech and language development low
IQ scores. - Usually caused by trisomy 21 (extra copy of a
chromosome 21). - During embryo formation, parents (usually
mothers) chromosome fails to split properly.
Laura Dwight
73Brain Abnormalities and Memory Impairments
- In Down syndrome, brain size may be average at
birth, but growth in some areas (e.g.,
hippocampus, frontal cortex, cerebellum) may be
stunted. - Individuals tend to have profound deficits in
hippocampal-dependent memory abilities. - Young adults with Down syndrome performed at
the 5-year-old level on mental abilities tasks. - Also, performed much worse on
hippocampal-dependent memory tasks.
74Hippocampal-DependentLearning and Down Syndrome
Data from Vicari, Bellucci, Carlesimo, 2000).
75Brain Abnormalities and Specific Memory
Impairments in Down Syndrome
Figure summarizes performance on battery of tests
that require hippocampal function (like list
learning and spatial learning) compared with a
battery of tests that require prefrontal function
(like working memory). Adapted from Pennington
et al., 2003, Figure 2.
76Animal Models of Down Syndrome
- Mice bred for segmental trisomy (Ts65Dn mice)
showed deficits in hippocampal-dependent tasks
(e.g., location of maze goal). - Enriched environment improved spatial memory in
female Ts65Dn mice. - Exacerbates impairment in Ts65Dn males.
77Alzheimers Disease
- Alzheimers Disease (AD)a form of progressive
cognitive decline from accumulating brain
deterioration. - AD affects about 4.5 million people in U.S.
- As many as 50 percent of people over age 85 are
afflicted. - http//www.youtube.com/watch?v7-P9lbTJ9Hw
78Progressive Memory Loss and Cognitive
Deterioration
- AD progression
- Earliest symptoms of AD occur in episodic
memory, such as forgetting recent visitors. - Later, there are declines in semantic memory
(e.g., forgetting familiar names, locations). - Next, conditioning and skill memory
deteriorate. - In late-stage AD, there is often a lack of
awareness and daily living skills. - http//www.youtube.com/watch?voTEbq4h-kvQ
79Patients with AD show marked impairment in many
forms of memory, including list learning. Over
three trials with a 10-word list, AD patients
recall fewer items than same-aged healthy
controls after a 10 minute delay, the patients
recall almost none of the studied words. Adapted
from Figure 1 of Moulin et al. (2004).
80Plaques and Tangles in the Brain
- Amyloid plaques deposits of beta-amyloid
(abnormal byproduct of amyloid precursor protein,
or APP kills adjacent neurons). - Plaques are fairly evenly distributed across
cerebral cortex. - Neurofibrillary tangles collapsed protein
scaffolding within neurons. - Early in AD, accumulate in hippocampus and MTL,
relating to semantic and episodic memory
deficits. - Hippocampal shrinkage early AD warning sign.
81Plaques and TanglesHallmarks of Alzheimers
Disease
a) Amyloid plaque (dark center spot) surrounded
by residue of degenerating cells.
b) Neurofibrillary tangles (seen as darkened
areas).
(a) Cecil Fox/Science Source/ Photo Researchers.
(b) Adapted from Figure 3 of Hardy Gwinn-Hardy,
1998.
82Plaques and Tangles in the Brain
- Verification of presence of plaques and tangles
(to confirm AD diagnosis) can only happen at
autopsy. - 10 to 20 percent of probable AD diagnoses
(based on MRI, PET, lumbar puncture, etc.) are
incorrect. - Many other conditions (some treatable) mimic
AD, so better diagnostic test needed. - e.g., vitamin B deficiency, hypothyroidism,
depression
83Genetic Basis of Alzheimers Disease
- Several genes implicated in AD.
- Most progress understanding genetic cause of
early-onset AD (begins at 3550 years). - Less than 1 percent of AD cases early-onset.
- Caused by genetic mutations, which are
autosomal dominant (meaning, just one mutated
gene from either parent will trigger early-onset
AD).
84Connection Between Down Syndrome and Alzheimers
Disease?
- Chromosome 21 (implicated in Down syndrome)
contains APP (implicated in AD). - By age 3540, adults with Down syndrome develop
neural plaques and tangles. - Half of Down syndrome patients show memory
decline and other symptoms of AD other half do
NOT show cognitive decline. - Why? Unclear. Explanation will help in
understanding both pathologies.
85Unsolved MysteriesTreating (and Preventing)
Alzheimers Disease
- Cholinesterase inhibitors treat forgetfulness and
anxiety. - Inhibiting breakdown of neurotransmitter
acetylcholine (depleted in patients with AD). - Memantine blocks glutamate receptors.
- May help protect neurons from
glutamate-mediated damage, slow cognitive decline.
86Unsolved MysteriesTreating (and Preventing)
Alzheimers Disease
- Risk factors for AD include
- Type-II diabetes
- High LDL (bad cholesterol)
- Previous head injury
- Stroke
- High blood pressure
- High levels of cognitive activity may slow AD
symptoms.
8712.3 Interim Summary
- Down syndrome condition in babies born with
extra copy of chromosome 21. - Children with Down syndrome have cognitive
impairments. - Includes memory impairments.
- Some brain areas tend to be abnormally small.
- Includes hippocampus, frontal cortex,
cerebellum.
8812.3 Interim Summary
- In Alzheimers disease, plaques and tangles
accumulate in the brain. - Memory symptoms are prominent early in the
disease. - Consistent with finding that hippocampus and
nearby MTL areas suffer pathology early in
disease. - Several genes may contribute to an individuals
risk for the common, late-onset form of the
disease.