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Figure 25. Simple apparatus built with a in Neurobiology and vision of jumping spiders (Araneae: Salticidae)
Figure 25. Simple apparatus built with a balloon to represent the eye tube of the right AME, a roll of paper to secure the anterior end of the tube, and attached rubber bands used to model the action of each of the four (1-4) extraocular muscles, as well as the two circular ocular muscles (5, 6). Muscle numbers correspond to those assigned by Land (1969b). By pulling on one or more rubber bands, the likely action of the respective muscles could be observed directly. The action of the circular muscles was more challenging to model, but I was able to simulate their contraction by pulling the ends against a loop in the middle of each band.
Figure 26. Four frames from a in Neurobiology and vision of jumping spiders (Araneae: Salticidae)
Figure 26. Four frames from a video of an adult female Colonus sylvanus in Greenville County, South Carolina. The transparency of the prosoma allowed a direct view of the movements of the AME. 1, This spider faced a Leucauge venusta (Araneae: Tetragnathidae) suspended under a nearby grass blade. The axis of the prosoma (blue line) and the optical axes of the AME (red lines) faced the prey directly. 2, The eye tube of the right AME moved to the left, shifting the axis of that eye to face the stem that would serve as an indirect route of access to that prey. 3-4, The spider slowly turned to the right, lowering its profile in the prey direction (upper left), and faced that access route. Subsequently, this Colonus slowly climbed the nearby stem, then moved under the attached grass blade to approach its prey, which was then captured with an upside-down jump. Active movement of the two AME tends to be loosely coupled, but each eye can also be moved independently a shown in (2).
Figure 24 in Neurobiology and vision of jumping spiders (Araneae: Salticidae)
Figure 24. Schematic frontal view of the arrangement of muscles associated with the AME, after Land (1969b, Pelegrina aeneola). Land described how, in dissection, these appeared as two sets per eye (1-6-3 and 2-5-4). Muscles 1-4, originating on the carapace, are extraocular muscles. Note that the dorsal muscles (3, 4) originate toward the rear of the ocular quadrangle, and the ventral muscles (1, 2) originate at the clypeus. The wide but thin circular muscles (5 and 6) are ocular muscles. These encircle the eye tube obliquely on either side, joining the extraocular muscles where they originate at either the top or the bottom of the eye tube. Muscle 2 divides into two branches, one of which (2b) joins muscle 1 before a common lateral origination point on the clypeus.
Figure 17 in Neurobiology and vision of jumping spiders (Araneae: Salticidae)
Figure 17. Stained sections through the AME and associated visual centers of Phidippus. 1, Horizontal section through prosoma of an adult female P. clarus. 2, Another horizontal section of the right AME and AME optic nerve of an adult female P. clarus. 3, Detail from (2), showing receptor tiers in the fovea. 4, Detail of AM1 in horizontal section, close to (1). 5, Parasagittal section of prosoma of fifth instar male P. clarus, showing dorsal and ventral lobes of the AME retina. 6, Transverse (frontal) section through prosoma of sixth instar P. johnsoni. 7, Transverse section through prosoma of a sixth instar P. johnsoni, through the boomerang-shaped retinae of the AME.
Figure 22 in Neurobiology and vision of jumping spiders (Araneae: Salticidae)
Figure 22. Relationship of tiered AME receptors to the first AME neuropile (AM1). 1, Camera lucida drawing of a Golgi- Kopsch impregnation of two neurons of the left AME (sixth instar Phidippus johnsoni), suggesting that the relative position of receptors in the retina correponds to the relative (retinotopic) position of corresponding, columnar terminals in the cortex of AM1. 2, Semi-diagrammatic drawing of a section through the inner (at left) and outer (at right) lobes of a Phidippus AM2, showing a 1:1 correspondence between columnar terminals of the two layers (after Hill 1975, 2006; Oberdorfer 1977). 3, Diagrammatic antero-dorsal view of the left AM1 of Phidippus. Fibers and terminals of the upper outer lobe are larger than those of the inner lobe and the lower part of the outer lobe. 4, Three views of the inner (yellow) and outer (green) lobes of the AM1. 5, Mapping of tiered AME receptors (I-IV) to respective terminal zones (TZ1-TZ4) of the AM1 of Hasarius adansoni (after Nagata, Arikawa & Kinoshita 2019). 1-4, after Hill (1975, 2006).
Figure 6 in Neurobiology and vision of jumping spiders (Araneae: Salticidae)
Figure 6. Camera lucida drawings of 10 μm sections of the prosoma of a fifth instar Phidippus clarus. 1, Parasagittal section. Visual centers of the AME (AM1, AM2) lie at the top of the protocerebrum. Each corpus pedunculatus lies beneath the AME neuropiles, and forms compact synapses (glomeruli of the lame glomerulee) with the ipsilateral lateral eyes (ALE and PLE). A lateral esophageal dilator separates the cheliceral and pedipalpal ganglia on each side. The dorsal and ventral lobes of the central body are situated at the rear of the protocerebrum. Densely packed neuronal cell bodies are peripheral to the mass of the CNS. In this vertical section, only the dorsal and ventral extensions of layers I and II of the retina of the AME can be seen. 2, Sagittal (midsagittal) section. The rostral ganglion, situated above the anterior esophagus, sends the recurrent nerve to the rear just above the esophagus, and the rostral nerve to the front. Commissures of the respective segmental ganglia are separated by blood vessels, associated with tracheoles that originate with the opisomal tracheal spiracle. These tracheoles pass between the sucking stomach and the mass of fused segmental ganglia, then turn ventrally to penetrate that mass at the midline. After Hill (1975, 2006).
Figure 16 in Neurobiology and vision of jumping spiders (Araneae: Salticidae)
Figure 16. General structure of the AME and related visual centers. 1, Camera lucida drawing of a parasagittal section of the anterior prosoma of a fifth instar Phidippus clarus. Since retinal layers I and II are dorso-ventrally elongated and boomerangshaped, only the dorsal and ventral arms of these layers can be seen in this view. The columnar matrix that is shown here is quite visible in stained sections, but has not been mentioned in other studies. In older spiders, the midgut digestive diverticulae (dd) are much larger than this. 2, Camera lucida drawing composited from several horizontal sections of the prosoma of a sixth instar P. johnsoni. In this view only the central, foveal part of the retina can be seen. Sections of the outer and inner lobes of the first AME neuropile (AME I) can be seen. 3, Semi-schematic dorsal view with the AME of a fifth instar P. clarus presented in horizontal section, showing the relationship of the eyes to the reconstructed lobes of the AME I neuropile. As a convention with the AME and the other eyes, distal refers to a direction along the optical axis toward the cornea, and proximal is a direction away from the cornea, toward the retina and optic nerve. As noted by Land (1969a), each AME optic nerve rotates by ~90° between the eye and AME1, although the degree of this rotation varies when the eye-tube is rotated. Note that some shrinkage and distortion did occur as part of the preparation of these sections, which were stained with Masson Trichrome. After Hill (1975, 2006).
Figure 19 in Neurobiology and vision of jumping spiders (Araneae: Salticidae)
Figure 19. Arrangement of bipolar receptor neurons in the AME of Pelegrina aeneola (Curtis 1892), after Land 1969a, 1985b. 1, Semi-schematic drawing of horizontal section through the proximal end of the AME. 2, Plotted position of receptors for each layer (I-IV) of the retina (anterior projection). Blue rectangles occupy the same position in a parasagittal (axial) projection.
Figure 21 in Neurobiology and vision of jumping spiders (Araneae: Salticidae)
Figure 21. Schematic cross sections of the AME retina (layer I). All are packed in a space-filling hexagonal array, although the paired rhabdoms of Goleba puella (Simon 1885) are rectangular (1). One receptor cell is highlighted in green in each image. The diameter of each receptor varies, with the smallest receptors, more closely packed, near the center of the retina. Each receptor is surrounded by the projections of unpigmented glial cells. 1, Goleba puella, after Blest, O'Carroll & Carter 1990. 2, Portia fimbriata (Doleschall 1859), Spartaeus spinimanus (Thorell 1878), or immature Servaea vestita, after Blest & Price 1984); Blest & Sigmund 1984); Blest 1985b; Blest & Carter 1987, 1988. 3, Lyssomanes viridis (Walckenaer 1837), Cyrba algerina (Lucas 1846), Spartaeus spinimanus, or Yaginumanis sexdentatus (Yaginuma 1967) after Blest & Sigmund 1984, 1985; Blest, O'Carroll & Carter 1990. 4, Colonus sylvanus or immature Servaea vestita, after Blest & Carter 1987, 1988; Blest, O'Carroll & Carter 1990. 5, Phidippus, Cyrba algerina,?Jacksonoides kochi (Simon 1900) or Lyssomanes dissimilis Banks 1929, after Eakin & Brandenburger 1971; Blest & Price 1984; Blest & Sigmund 1984; Blest, McIntyre & Carter 1988; Blest, O'Carroll & Carter 1990. 6, Phidippus johnsoni, lateral, after Eakin & Brandenburger 1971. Note reduced packing and paired rhabdoms of these larger, lateral receptors. In some species the presence of one or two rhabdom groups in each cell depends on the plane of section.
Figure 5 in Neurobiology and vision of jumping spiders (Araneae: Salticidae)
Figure 5. Camera lucida drawings of half transverse (frontal or cross) 10 μm sections of the prosoma of a fifth instar Phidippus clarus (1), and a sixth instar P. johnsoni Peckham & Peckham 1883 (2). In each drawing the midline is at the right. 1, Section through the posterior lateral eye (PLE). The short optic nerve of each lateral eye (ALE and PLE) has its own folded first neuropile (PL1 for the PLE). interneurons associated with this neuropile are associated with tracts leading to both the glomerlar synapses (or glomeruli) of the corpora pedunculata, and to a lateral eye neuropile. 2, Section through the sucking stomach. The CNS is separated from the sucking stomach by the endosternite, an internal skeletal element comprised of a cartilage-like material. This view emphasizes the powerful musculature that fills much of the prosoma. Many of these striated muscles are attached to the endosternite. After Hill (1975, 2006).
Figure 2 in Neurobiology and vision of jumping spiders (Araneae: Salticidae)
Figure 2. Semi-schematic views of the CNS exposed in a dissection of the prosoma of Phidippus jumping spiders. 1, Adult female Phidippus clarus Keyserling 1885. The optic nerves of the anterior medial eyes (AME) lead directly to the rear, to the AME neuropiles I and II at the top of the CNS. Below these are the wide optic nerves of the anterior and posterior lateral eyes (ALE and PLE, respectively), and the small optic nerves of the posterior medial eyes (PME). In most salticids the PME are small as shown here, but in some they are much larger. The fused opisomal ganglia (CE, or cauda equina) lie at the rear of a suboral, ventral mass of fused pedipalp and leg ganglia. 2, Visual fields of the respective eyes. The PLE have wide fields of vision, the ALE much narrower fields that overlap in front of the spider. The tubular AME have narrow fields, but are moved up and down and side to side to scan a larger field. After Hill (1975, 2006).
Figure 13 in Neurobiology and vision of jumping spiders (Araneae: Salticidae)
Figure 13. Thick (80 μm) sections of Golgi-Kopsch preparations of the prosoma of sixth instar Phidippus johnsoni. 1, Parasagittal section. 2,4, Horizontal sections. 3, Sagittal (midline) section. Golgi-Kopsch preparations can selectively stain nerve fibers, but it is not possible to determine just what fibers will be stained, and the results can be misleading. Here some of the major fiber tracts in the subesophageal part of the CNS have been stained.
Figure 1 in Neurobiology and vision of jumping spiders (Araneae: Salticidae)
Figure 1. Faces of two jumping spiders of the North American genus Phidippus C. L. Koch 1846. 1, Adult male P. pacosauritus Edwards 2020, Paco's Reserva de Flora y Fauna, MazatlaDn, Sinaloa, Mexico. 2, Adult female P. putnami (Peckham & Peckham 1883), southern Greenville County, South Carolina, USA.
Figure 9 in Neurobiology and vision of jumping spiders (Araneae: Salticidae)
Figure 9. Sections through the CNS, stained with reduced silver impregnation to highlight nerve fibers. 1, Half transverse section between the first and second legs of a sixth instar Phidippus johnsoni. Longitudinal tracts (in a plane perpendicular to the page) are highlighted in yellow. At multiple levels commissures, or transverse tracts cross the midline, connecting the right and left ganglia of each segment. The CP tract originates with the posterior end of a corpus pendunculatus and the ipsilateral lateral neuropile of the protocerebrum. 2, Sagittal section through the fused subesophageal ganglia of a fifth instar P. clarus, showing medial longitudinal nerve tracts and segmental commissures separated by blood vessels. A recurrent nerve runs along the top of the esophagus, just behind the preoral rostral ganglion.
Figure 20 in Neurobiology and vision of jumping spiders (Araneae: Salticidae)
Figure 20. Arrangement of receptors in the tiered retina of the AME. 1, Anterior view of an adult female Colonus sylvanus (Hentz 1846), depicting the boomerang shape of the AME retinae (in green), and the alignment of respective fields of vision (outlined in yellow). White arrows depict the six directions in which each AME can be moved, to include left-right, up-down, and clockwise-counterclockwise rotation. Each of these directions corresponds to the contraction of a set of oculomotor muscles that position the long AME eye tube within the prosoma. After Hill 2018a. 2, Parasagittal diagram of the respective alignment of receptors in the four layers of the AME of Servaea vestita, near the center of the retina. The identity of the receptors shown in orange was not known. Red rectangles show the position of columns connecting layers I and 2. After Blest et al. 1981. 3-6, Diagrams of serial (proximal to distal) transverse sections through the retina of S. vestita, showing a cross section of individual receptors by layer (after Blest et al. 1981; Blest 1988). Lateral receptors of layers III-IV include groups 3a, 4a and 4c. Medial receptors of these layers include group 3b and 4b. Based on the findings of Nagata, Arikawa & Kinoshita (2019; see Figure 22:5), the separation of layer III and IV receptors into these lateral and medial groups may have more anatomic or functional significance than their relative position (or layer) along the axis of the eye.
Figure 8 in Neurobiology and vision of jumping spiders (Araneae: Salticidae)
Figure 8 (continued on next page). Semi-schematic camera lucida drawings of a series of 10 μm half-horizontal sections through the prosoma of a fifth instar Phidippus johnsoni. In each drawing the midline or sagittal plane is shown at right. Numbers in rectangles (1-102) indicate the relative vertical position of each section in the series. These drawings illustrates the relative position of the CNS at various levels. Musculature in the plane of each section is shown in red, digestive diverticulae of the midgut in yellow. Nuclei of the unipolar neurons that surround the otherwise fibrous nerve masses are shown as small circles.
Figure 18 in Neurobiology and vision of jumping spiders (Araneae: Salticidae)
Figure 18. Oblique horizontal section through the anterior prosoma of a second instar Phidippus johnsoni. This was a 5 μm Epon section stained with Toluidine Blue, and thus very little shrinkage of structures occurred. Note the smooth pit and staircased (placed in multiple focal planes) receptors of layers I and II in the foveae of the AME, at center. The intrusive pigment at the margins of each AME eye tube is aligned with thin, membraneous structures within the clear matrix. Receptor layer I is most distinct, and nothing can be seen of receptor layers III and IV in this view.
Figure 7 in Neurobiology and vision of jumping spiders (Araneae: Salticidae)
Figure 7. Camera lucida drawings of parasagittal sections through the prosoma of Phidippus clarus at three different stages of development. Relative to body size, the CNS of the adult is relatively small. The prosoma of the adult is packed with midgut digestive diverticulae (highlighted in orange) above, below, and in front of the CNS. In the adult female, these supplement the diverticulae of the opisoma, as developing ova occupy an increasing portion of the opisomal volume. Recent studies have shown that adult and juvenile salticids have similar numbers of visual receptor cells, more tightly packed in the latter (GoteDet al. 2019). After Hill (1975, 2006).
Figure 15 in Neurobiology and vision of jumping spiders (Araneae: Salticidae)
Figure 15. Hypothetical wiring diagram depicting the connectivity of primary visual centers in the CNS of salticine jumping spiders. Based on studies of Hasarius adansoni (Nagata, Arikawa & Kinoshita 2019), Marpissa muscosa (Steinhoff et al. 2017, 2020), Phidippus (Hill 1975, 2006; Long 2016), Evarcha arcuata/Servaea vestita (Duelli 1980), and Salticus scenicus/Naphrys pulex (Oberdorfer 1977). In salticines, the unipolar receptor somata of the secondary eyes (ALE, PLE, with the possible exception of the PME) are located just outside of the respective eye cup as depicted here (Maddison & Hedin 2003). These synapse with interneurons in the highly convoluted cortex of the AL1 and PL2, and a separate, small PM1 neuropile. Secondary fibers of unipolar neurons with somata in the cortex of the CNS connect these to either the glomerular synapses (lame glomerulee) of the corpora pedunculata (with separate but adjacent AL2 and PL2 lobes as shown here), or to a less structured lateral neuropile (lenp, lateral eye neuropile). The bipolar sensory cells surrounding the retinal layers of the AME send a distal receptor process into the retina, and a proximal axon within the AME optic nerve, terminating in the synapses of the highly-structured AM1. Interneurons connect these to a less-structured, posterior neuropile (AM2). Heavy vertical bars at left indicate where fiber tracts continue into other parts of the CNS. Small neuropiles (PLx and ALx) connecting receptors of the respective lateral eyes to a lateral fiber tract may be either primary (as depicted here) or secondary. The PLx neuropile has only been documented in Marpissa (Steinhoff et al. 2017, 2019).
Figure 4 in Neurobiology and vision of jumping spiders (Araneae: Salticidae)
Figure 4. Camera lucida drawing of a parasagittal section through the prosoma of a second instar (emergent) Phidippus clarus. 1, At this stage, the CNS occupies nearly the entire medial volume of the prosoma. 2, Detail of inset from (1) showing a cluster of large neuron cell bodies in the cortex of the third and fourth leg ganglia, surrounded by smaller neurons. After Hill (1975, 2006).
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Allen Brain Atlas
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DANDI Archive for NWB datasets
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International Brain Laboratory public data
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OpenNeuro
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