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Pine Siskin flocks are constantly atwitter with wheezy contact calls while feeding or in flight. Their most recognizable call is a "watch-winding" note, a harsh, upsweeping zreeeeeeet lasting most of a second, tossed in amidst shorter calls. They utter a distinctive flight call, an explosive zwee or psee that initiates startled flight. In flight, a tit-a-tit call often accompanies each flap-and-glide undulation. A single siskin can call back the flock with a solitary note. Females solicit copulation by uttering soft calls while bowing and fluttering tail and wing feathers. The female solicits feeding from the male with a low twittering call that carries well through the trees.

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Sometimes it feels wrong to use the word "song" for an instrumental piece of music. You wouldn't call Miles Davis' "So What?" a song.* What would you call it? A "recording" works in some contexts, but not when referring to live music or to the [song] in general. "Piece of music" is awkward - in some contexts more than others if feels artificial, like there ought to be a better word. What word?

Or you can refer to a minuet or some specific type of [song]. If however you had a set list including a minuet and a tarantella, what would you call the items on that list? You wouldn't call them all minuets. Would you call them all songs?

"Call on Me" is a song written by Lee Loughnane for the group Chicago and recorded for their album Chicago VII (1974). Peter Cetera sang lead vocals and the arrangement makes prominent use of conga drums played by Guille Garcia.

"Call on Me" was the first Loughnane composition to be released by the band. It was written with uncredited help from Peter Cetera.[1] Loughnane was the last original Chicago member to receive a songwriting credit.

According to Cetera, though, he needed some help. "I tried to help Lee Loughnane with a song," Cetera says, "and that song turned out to be 'Call On Me.' Lee had written a song. It wasn't called, 'Call On Me,' it was called something else, and it in fact was terrible. I talked to him at the ranch one day, and he was all bent out of shape. He said that he had played the song for the guys, and they had told him in fact to get the heck out of there with the song. I said, 'Well, come on, let's have a go.' So Lee and I went and re-wrote the lyrics and re-wrote the melody and came up with the song called, 'Call On Me,' which was a big hit for him." Loughnane remembers it a little differently. "Peter changed a couple of the words and the way he sang the melody in order for him to be able to play the bass and sing the melody at the same time because that's the way he felt it." Loughnane added, "I appreciate his efforts, and we did make the song a hit."

Songbirds, which make up about half of all extant bird species, have the ability to learn complex vocalizations like song and certain types of distance calls beside their innate call repertoire, whereas the closely related suboscine species produce only unlearned song and calls. The emergence of the ability to produce learned vocalizations is associated with the evolution of the forebrain vocal control system, an interconnected network of brain nuclei that shapes the song during learning and organizes the motor output when singing [1], [2]. At best, only rudimentary traces of this system are found in the non-learning relatives of songbirds [3]. Therefore, the vocal control system is thought to be uniquely devoted to the control of learned sounds. The evolutionary steps that led to the development of the learning-related forebrain vocal system are as yet unknown, but it seems reasonable to assume that the song system has evolved from circuits driving simpler unlearned vocalizations.

Since, like stack calls, song is used in a social context, the association of the song control system with communicative calling activity might shed light on its evolutionary history. The premotor nucleus RA (nucleus robustus arcopallialis) is part of the song motor pathway [1], [9]. RA is electrically active during the production of learned vocalizations [10], [11] and controls the spectral and temporal properties of song elements of zebra finches [12]. RA, therefore, is a logical starting point for associating brain activity with stack calling exchanges. Moreover earlier studies suggest an involvement of RA in unlearned call production [12], [13]. Therefore, we studied brain and auditory activity in small groups of socially interacting zebra finches, each animal carrying a wireless microphone. Each male had electrodes implanted into RA to record neuronal signals during vocal production while moving freely. In this way we show that neurons in the song control system perform precise and pronounced burst firing prior to stack calling. Thus, RA has a function in the control of unlearned vocal social interactions. Based on this, we propose an evolutionary scenario in which the song control system evolved from a system that controlled unlearned sounds that were used to communicate with particular conspecifics in a social group; a process that involves learned sensory-motor integration.

The production of the stack calls by two partners was clearly time-locked, indicating that many calls were produced in response to the calling of the partner (Figure 2A). Out of the 35 pairs recorded, only two pairs did not show clear correspondences between stacks. The number of stacks that were themselves an answer, and the number that was answered showed less variation than the total numbers produced: in each of 5 pairs that we analyzed in detail and had developed a symmetrical calling relationship the number of calls produced varied 7-fold, whereas the number of stacks that were either an answer or received an answer varied between 1320 and 558, slightly more than twofold (Figure 2B).

In 20 pairs that were kept in soundboxes and recorded with a central microphone, the males carried a chronically implanted tungsten electrode connected to a transmitting high-impedance amplifier [14] to record electrical activity in RA while free moving. In all cases we could differentiate between two different stack calls that were, in 17 cases exchanged between partners (Figure S10). One of these stacks was invariably associated with RA activity (Figure 4; Figure S10). This stack call was in all probability the call of the male. Moreover, in 5 experiments where the stack calls could be attributed unequivocally to each of the individuals through the use of backpack microphones, the male call was always associated with RA activity (Figure S11).

The associated firing patterns in RA in 26 units from 25 birds can be classified as follows: Excitation before and during the call occurred in 15 cases, there were 6 instances of inhibition, a biphasic response (inhibition followed by excitation) occurred in 2 cases. Three units showed no significant response. Figures S10 and S11 summarize the results for all RA recordings. We have observed no instance where a stack call was incorporated into a song motif, even though many songs contain syllables that have a stacked sonogram.

In sum, these results show not only that RA firing is associated with stack calling, but also that the same units may be involved in the production of both the unlearned stack call and the learned song.

Since pair-bonded zebra finches live in larger social groups the observed specific call exchanges among group members require to learn the individual call signature of other group members and to respond in a timed fashion to specific calls. Such timed responses are not an automatism since we find that not all calls of the mate are answered or are answers. Short-range contact calling could help the animals to locate their partners in a flock. Why not all stack calls of the mate are answered despite being uttered within hearing range of the call receiver remains unknown.

The fact that RA is controlling innate calls as well as learned vocalizations allows speculation about the evolutionary origin of the song control system. Since vocal learning occurs only in three not closely related avian taxa (songbirds, hummingbirds, parrots) and since the closest relatives of the songbirds, the sub-oscine passerines do not show vocal motor learning [23], it is parsimonious to assume that production of innate sounds is the evolutionary older situation.

RA is most clearly defined both morphologically and neurophysiologically in oscine songbirds. In suboscines areas analogous to RA have been described [3], and show concentrations of RA-like cells, but this cell group is not nearly as clearly delineated as in songbirds. During songbird evolution, this RA-precursor could have extended its role from the control of innate calls to learned songs. Our finding that many RA neurons fire milliseconds before innate calls are produced, supports this hypothesis. In particular, since stack call controlling neurons are also involved in the production of learned song syllables, RA is not composed of two separate sub-circuits dedicated to either learned or innate sounds but the same neurons do both, i.e. carry out an evolutionary basic and a derived task. Further, the RA firing patterns suggest an involvement not only in calling per se but also in precisely timed call exchanges between partners, which requires learning.

The study of brain activity in awake birds has contributed greatly to our understanding of bird song learning and production [10], [29], [30]. Until now, recording in moving animals imposed restrictions on the freedom of movement, because the animals were tethered, which made it impossible to study neuronal activity in social groups. With our lightweight radio transmitters we make available a method that allows us to record the signal of deep brain electrodes and individual vocalizations synchronously. This enables us to relate individual signaling behavior with the underlying neuronal pattern in a group of zebra finches living in an aviary which provides insight into the evolutionary link between innate call production and learned song. e24fc04721