The Numerate Brain
What is Number?
Number can be a symbol, a unit of measurement, a quantity, a code, or a magnitude.
How do we study number ability?
Relative vs. Absolute judgments.
Symbolic vs. Non-symbolic representations.
Subitizing as a special case.
Is our sense of number innate and unique?
Where there is ability, there is a biological process.
Number in the brain.
Spatial representations.
Models of Number Processing
Cognitive models and how they deal with different representations and/or transcoding (switching forms).
Visual Number Perception
Factors that influence judgments of visual quantity/magnitude.
2D vs. 3D.
Visual Models
Is number coded or is it the density of items in an area?
The primacy of our number sense.
Dakin’s model for coding number per area (i.e., coding number as density of items).
The Meaning of Numbers
The meaning of a number is a quantity, magnitude, or size of a collection (numerosity).
A telephone number is a code that uses number symbols but doesn’t convey numerical meaning.
Meaning is abstract.
It is independent of the symbol denoting it (3, three, trois, 3 fingers) and the object being enumerated (3 ideas, 3 elephants, 3 computers).
Non-Symbolic Number Cognition: Quantities
Tasks often involve judging the size of an array.
Which has more? (relative).
Are there 8? (absolute).
Relative judgments of array size depend on:
Number of items (size effect) e.g. 20:10 harder than 10:5.
Difference in set size (distance effect) e.g. 6:5 harder than 7:5.
Performance correlates with school-based maths measures (Halberda et al. 2008).
Impaired in developmental dyscalculia (Piazza et al. 2010).
Quantities
Absolute judgments of array size (exactly how many?) depend on two mechanisms:
Subitizing (up to 3-4 items at a glance, highly accurate, parallel processing).
Counting (4+ items, conversion to symbolic code, serial processing, error-prone).
Approximating- we don’t/can’t always count.
Symbolic Number Cognition
Deciding which of two number symbols is larger depends on:
The distance between the numbers (distance effect; e.g. "5 4" slower than "5 3").
The size of the numbers themselves (size effect; e.g. "5 3" slower that "4 2").
Interactions Between Symbolic & Non-Symbolic Number Codes
Congruent Examples:
Which is physically larger?
2 (423 msec) vs. 9 (532 ms).
Which is numerically larger?
2 vs. 9.
Incongruent Examples:
Which is physically larger?
9 vs. 2.
Which is numerically larger?
2 (450 msec) vs. 9 (619 msec).
Number size influences physical size judgment.
Physical size influences number size judgment.
Girelli, Lucangeli and Butterworth (2000).
Interactions Between Symbolic & Non-Symbolic Number Codes
Koechlin et al. (1998).
Priming study:
Judge whether a stimulus is greater than 5.
Stimuli consist of names (SEVEN), digit (7) or dot arrays.
Before each trial, participants are shown a brief subliminal prime that is either on the same side of 5 as the target, or the opposite side.
Interference is found when the prime and target are on opposite sides of 5.
Influences extend across representations.
Suggests semantic priming.
Number Sense
Ability to sense, compare, and estimate numbers.
How common is this?
How innate is this?
How does the brain do it?
Innate Versus Cultural Aspects of Number
Names within the subitizing range seem to be a general ability across cultures (Amazonian Munduruku number naming).
Implies a common underlying mechanism or set of mechanisms.
Other aspects of numerical cognition (such as written symbols, number names, equations, algebra etc.) may be cultural inventions.
An appreciation of quantity (how much?) and numerosity (how many?) may be a basic skill of almost all humans and adults.
Infants habituate if shown the same number of items but dishabituate if the number changes.
Number Neurons?
Animal work: physiology measuring action potentials (Nieder, 2005).
Some respond to sequential presentation of dots as well as simultaneous ones.
Some respond to the number of sounds too (i.e. multi-sensory coding of number).
Training monkeys to associate digits to set sizes reveals neurons (in PFC, not IPS) that respond to symbolic and non-symbolic number.
Human children show frontal-to-parietal development shift in processing number (fMRI); suggesting symbols become linked to ‘core’ number code of meaning in IPS over time.
A Neural Region For Number Meaning?
Converging evidence suggests the importance of a region in the intraparietal sulcus (IPS).
More active in calculation than number reading.
Sensitive to subliminal priming when prime and target differ in number.
Shows a distance effect for both digits and number words.
Habituation to repeated numbers of items in a dot array.
A Separate Region for Visual Recognition of Number Symbols?
Left hemisphere.
Preference for shapes.
Biased connectivity to language.
Right hemisphere.
Biased connectivity to numerosity.
T&F, NFA, VWFA.
Left and Right Brain in Number Meaning
Studies of acquired/developmental dyscalculia emphasize the importance of the left hemisphere, but functional imaging studies show bilateral activity. Why?
One suggestion is that the left hemisphere deals with exact quantities (e.g., 5), whereas the right hemisphere deals with approximate quantities (e.g., between 4 and 6).
Warrington (1982): dyscalculic patient with LH lesion able to do approximate sums (e.g. = “roughly 13”).
Dehaene and Cohen (1996): digits presented to LH and RH of a split-brain patient; RH gives approximate answers 5 "six" but LH gives exact answers.
Representing Number
Spatial Code for Numbers.
SNARC = Spatial numerical association of response codes.
The SNARC effect = faster response by left hand to small numbers, and right hand to large numbers.
Counting explicitly using body parts: The “Yupno” in Papua New Guinea and Torres Straight Islanders (Adapted from Ifrah (1985)).
Modeling Number Ability
Conceptualize number representation through models:
Cognitive models.
Visual models.
Models of Numerical Cognition
McCloskey Model.
Cognitive model.
Number size is represented as base-10 units (divisible into 10s, 100s, 1000s, etc.).
Separate routines or stores for arithmetical operations ().
Abstract (semantic) representations are used for all calculations.
Transcoding is performed semantically.
Dehaene's Triple-Code Model.
Cognitive and neuroanatomical model.
Number size is represented in a logarithmically compressed form (larger numbers harder to discriminate).
No separate routines or stores for arithmetical operations ().
Some calculations are independent of number semantics (e.g., multiplication is verbal fact retrieval).
Transcoding may be performed without semantics.
How do interactions occur: Transcoding
Transcoding = translating one type of symbol (e.g., 5) to another (five); reading and spelling are types of transcoding.
Evidence suggests some non-semantic transcoding (favoring Dehaene’s model).
Cipolotti and Butterworth (1995) case study:
Asked to write "seventy thousand" he wrote 17,000 but when asked to do 56,748 + 13,252 he wrote 70,000.
Models of Numerical Cognition: A Hybrid Account
An adaptation of the McCloskey model by explicitly adding transcoding stages (From Butterworth (1999)).
How is number being encoded: Base-10 or Log-Linear
Some evidence for both theories.
McCloskey assumes separate representations for units, tens, hundreds, etc.
Evidence for this comes from e.g., deciding which digit is larger (e.g., 51 vs. 65), performance is affected by both tens (6>5) and unit magnitude (5>1) relative to when there is a mismatch.
Dehaene assumes a "mental number line" because number comparison shows the same law as comparing the length of two lines.
Models of numerosity perception
Visual models: how do we estimate non-symbolic quantities?
The previous models assume simple counting, but often we estimate numbers too rapidly to count (e.g., crowds, sheep).
When we do, we are reasonably accurate too (10-20% JND).
Professor John Ross did considerable work in this field.
Models of numerosity perception: Approximating number as density
We can estimate large numbers relatively accurately and fast without counting- but how?
Dakin et al, PANAS, 2011, argue that number is affected by area:
They showed that larger patches look more numerous (E), implying our sense of number is tied to the area being referenced.
Dots per area is saying we encode density rather than number…
But how to code: Approximating number as density
Larger patches look more numerous.
The Dakin model assumes that fine detail (High spatial frequency) processes the number of items.
By contrast, coarse detail (low spatial frequency) processes the relevant area.
Response: Number Sensed Directly
John Ross supported the Dahaene view that we sense number directly, i.e. number neuron/, logarithmic numberline.
John hated the number = density argument, and there were several exchanges in the published literature…
Durgin's suggestion, that number is extracted indirectly from a texture representation obtained from the statistical kurtosis of a scene evaluated over various scales, seems reminiscent of the legendary Australian stockman who, when asked to explain his uncanny ability to judge the number of cattle in a herd, replied that he counted the legs and divided by four.
In the end, as often happens, the answer seems to be that we use, or sense number for smaller N and use a density metric for larger N (e.g. crowds).
Anobile et al., Psychological Science, 2014.
Consistent with number neurons for smaller magnitudes.
Some of my work in this area: 2D or 3D
I’ve worked on the coding of number, testing Dakin’s model and whether it is right to be 2D or needs to incorporate 3D information.
2D or 3D
I replicated an area bias for density but NOT for number (argues against Dakin model).
I showed that density biases did not occur for 3D volumes.
How Primary is Our Sense of Number?
I’ve also looked at what types of visual cues interact with our sense of number: Apthorp & Bell, Current Biology 2015.
Symmetrical patterns appear less numerous (compare top row in A).
We argue this is because of redundancy. We need only attend and process half the pattern.
This biases our estimate of number down.
Implies symmetry is processed before number.
Summary
Number is a complex concept: symbols, magnitudes, codes, quantities.
There are several commonalities across these that point to a central abstract concept of number.
This concept seems to map to space (SNARC).
Number ability seems somewhat innate, crosses cultures.
Number involves several brain regions, visual, verbal, abstract number concepts.
These concepts relate to models of number.
The primacy of number has given rise to models.
Cognitive and visual.