跟著城市嚮導「老臺北胃」，用味道認識臺北

很多朋友來臺北，
都會問我同一個問題：
「臺北小吃那麼多，到底該從哪裡開始吃？」
夜市裡攤位一字排開、老店藏在巷弄轉角，
看起來都很有名，卻又怕吃錯、踩雷，
結果行程走完，反而沒真正記住臺北的味道。
我常被朋友笑說是「老臺北胃」。
不是因為特別會吃，而是因為在這座城市待久了，
知道哪些味道是陪著臺北人成長的日常。
這篇文章，就是我整理的一份清單。
如果你第一次來臺北，
我會帶你從這 10 樣最具代表性的臺北小吃開始，
不追一時爆紅、不走浮誇路線，
而是讓你吃完後能真正理解
原來，這就是臺灣的小吃文化。
跟著老臺北胃走，
用最簡單的方式，
把臺北的味道，一樣一樣記在心裡。

我怎麼選出這 10 大臺北小吃？

在臺北，
你隨便走進一條夜市或老街，
都可以輕易列出 30 種以上的小吃。
所以這份清單，
不是「臺北最好吃」的排名，
 而是我站在「第一次來臺北的旅客」角度，
做的推薦。
身為一個被朋友稱作「老臺北胃」的人，
我選這 10 樣小吃時，心裡一直放著幾個原則。

一吃就知道：這就是臺灣味

燒烤、火鍋很好吃，
但換個城市、換個國家，也吃得到。
我挑的，是那種
只要一入口，就會讓人聯想到的臺灣味。
 不需要解釋太多，舌頭就能懂。

不只是好吃，而是有「臺北日常感」

臺北的小吃迷人，
不只在味道，
而在它融入生活的方式。
我在意的是：

1. 會不會出現在早餐、宵夜、下班後
2. 有沒有陪伴這座城市很久的記憶

吃完之後，你會記得臺北

最後一個標準很簡單。
如果你回到家，
還會突然想起某個味道、某碗熱湯、某個攤位的香氣
那它就值得被放進這份清單裡。

接下來的 10 樣臺北小吃，
就是我會親自帶朋友去吃的在地美食。
不趕行程、不拚數量，
而是一口一口，
慢慢認識臺北。

第 1 家：饌堂－黑金滷肉飯（雙連店）｜一碗就懂臺灣人的日常

如果只能用一道料理，
 來解釋臺灣人的日常飲食，
 那我一定會先帶你吃滷肉飯。
在臺北，滷肉飯不是什麼特別的節慶料理，
 而是從早餐、午餐到宵夜，
 默默陪著很多人長大的味道。
而在眾多滷肉飯之中，
饌堂－黑金滷肉飯（雙連店），
 我很常帶第一次來臺北的朋友造訪的一家。

為什麼第一站，我會選饌堂？
饌堂的滷肉飯，走的是**「黑金系」路線**。
滷汁顏色深、香氣厚，
卻不死鹹、不油膩。
滷肉切得細緻，
肥肉入口即化，搭配熱騰騰的白飯，
每一口都是很完整、很臺灣的味道。
對第一次吃滷肉飯的旅客來說，
這種風味夠經典、也夠穩定，
不需要太多心理準備，就能理解為什麼臺灣人這麼愛它。

不只是好吃，而是「現在的臺北感」
饌堂並不是那種躲在深巷裡的老攤，
空間乾淨、節奏俐落，
卻沒有失去滷肉飯該有的靈魂。
這也是我會推薦給旅客的原因之一：
它保留了臺灣小吃的核心味道，
同時也讓第一次來臺北的人，
吃得安心、坐得舒服。

老臺北胃的帶路小提醒
如果是第一次來：

1. 一定要點招牌黑金滷肉飯
   
2. 可以加一顆滷蛋，風味會更完整
3. 搭配簡單的小菜，就很有臺灣家常感

這不是那種吃完會驚呼「哇！」的料理，
而是會讓你在幾口之後，
慢慢理解
原來，臺灣人的日常，就是這樣被一碗飯照顧著。

地址：103臺北市大同區雙連街55號1樓

電話：0225501379

菜單：https://bio.site/ZhuanTang

第 2 家：富宏牛肉麵｜臺北深夜也醒著的一碗熱湯

如果說滷肉飯代表的是臺灣人的日常，
 那牛肉麵，
 就是很多臺北人心中最有份量的一餐。
而在臺北提到牛肉麵，
 富宏牛肉麵，
 幾乎是夜貓族、加班族、外地旅客一定會被帶去的一站。

為什麼老臺北胃會帶你來吃富宏？
富宏最讓人印象深刻的，
不是華麗裝潢，
而是那鍋永遠冒著熱氣的紅燒湯頭。
湯色濃而不混，
帶著牛骨與醬香慢慢熬出的厚度，
喝起來溫潤、不刺激，
卻會在嘴裡留下很深的記憶點。
牛肉給得大方，
燉到軟嫩卻不鬆散，
搭配彈性十足的麵條，
每一口都很直接、很臺北。

不分時間，任何時候都適合的一碗麵
富宏牛肉麵最迷人的地方，
在於它陪伴了無數個臺北的夜晚。
不管是深夜下班、看完演唱會、
或是剛抵達臺北、還沒適應時差，
這裡總有一碗熱湯在等你。
對旅客來說，
這種不用算時間、不用擔心打烊的安心感，
本身就是一種臺北特色。

老臺北胃的帶路小提醒
第一次來富宏，我會這樣點：

1. 紅燒牛肉麵是首選
   
2. 如果想吃得更過癮，可以加點牛筋或牛肚
3. 湯先喝一口原味，再視情況調整辣度

這不是精緻料理，
卻是一碗能在任何時刻撐住你的牛肉麵。
在臺北，
很多夜晚，
就是靠這樣一碗熱湯走過來的。

地址：108臺北市萬華區洛陽街67號

電話：0223713028

菜單：https://www.facebook.com/pages/富宏牛肉麵-原建宏牛肉麵/

第 3 家：士林夜市・吉彖皮蛋涼麵｜臺北夏天最有記憶點的一口清爽

如果你在夏天來到臺北，
 一定會很快發現一件事
 這座城市，真的很熱。
也正因為這樣，
 臺北的小吃世界裡，
 才會出現像「涼麵」這樣的存在。
而在士林夜市，
 吉彖皮蛋涼麵，
 就是我很常帶旅客來吃的一家。

為什麼在夜市，我會帶你吃涼麵？
很多人對夜市的印象，
都是炸物、熱湯、重口味。
但真正的臺北夜市，
其實也很懂得照顧人的胃。
吉彖的涼麵，
冰涼的麵條拌上濃郁芝麻醬，
再加上切得細緻的皮蛋，
入口的第一瞬間，
就是一種「被降溫」的感覺。
那種清爽，
不是沒味道，
而是在濃香與清涼之間取得剛剛好的平衡。

皮蛋，是靈魂，也是臺灣味的關鍵
對很多外國旅客來說，
皮蛋是既好奇、又有點猶豫的存在。
但我常說，
如果要嘗試皮蛋，
涼麵是一個非常溫柔的起點。
芝麻醬的香氣會先接住味蕾，
皮蛋的風味則在後段慢慢出現，
不衝、不嗆，
反而多了一層深度。
很多人吃完後，
都會露出那種「原來是這樣啊」的表情。

老臺北胃的帶路小提醒
第一次點吉彖皮蛋涼麵，我會建議：

1. 一定要選皮蛋款，才吃得到特色
2. 醬料先拌勻，再吃，風味會更完整
3. 如果天氣真的很熱，這一碗會救你一整晚

這不是華麗的小吃，
卻非常臺北。
在悶熱的夜晚，
站在夜市人潮裡，
吃著一碗涼麵，
你會突然明白——

原來臺北的小吃，連氣候都一起考慮進去了。

地址：111臺北市士林區基河路114號

電話：0981014155

菜單：https://www.facebook.com/profile.php?id=100064238763064

第 4 家：胖老闆誠意肉粥｜臺北人深夜最踏實的一碗粥

如果你問我，
 臺北人在深夜、下班後，
 最容易感到被安慰的食物是什麼——
 我會毫不猶豫地說：肉粥。
而提到肉粥，
 胖老闆誠意肉粥，
 就是很多老臺北人口中的那一味。

為什麼這一碗粥，會被叫做「誠意」？
胖老闆的肉粥，看起來很簡單。
白粥、肉燥、配菜，
沒有華麗擺盤，也沒有複雜作法。
但真正坐下來吃，你會發現：
這碗粥，不敷衍任何一個細節。
粥體滑順、不稀薄，
肉燥香而不膩，
搭配各式家常小菜，
一口一口吃下去，
很自然就會放慢速度。
這種味道，
不是要你驚艷，
而是要你安心。

這不是觀光小吃，而是臺北人的生活片段
胖老闆誠意肉粥，
最迷人的地方，
就是它的客人。
你會看到：

1. 剛下班的上班族
2. 熬夜後來吃一碗熱粥的人
3. 熟門熟路、點菜不用看菜單的老客人

這些畫面，
比任何裝潢都更能說明這家店在臺北的位置。
對旅客來說，
這是一個走進臺北人日常的入口。

老臺北胃的帶路小提醒
第一次來吃，我會這樣建議：

1. 肉粥一定要點，這是主角
2. 配幾樣小菜一起吃，才有完整體驗
3. 不用急，慢慢吃，這碗粥就是要你放鬆

這不是為了拍照而存在的小吃，
而是那種
**會讓人記得「那天晚上，我在臺北吃了一碗很溫暖的粥」**的味道。

地址：10491臺北市中山區長春路89-3號

電話：0913806139

菜單：https://lin.ee/xxbYZyS

第 5 家：圓環邊蚵仔煎｜夜市裡最不能缺席的臺灣經典

如果要選一道
 最常出現在旅客記憶裡的臺灣小吃，
 蚵仔煎一定排得上前幾名。
而在臺北，
 圓環邊蚵仔煎，
 就是那種很多臺北人從小吃到大的存在。

為什麼蚵仔煎，這麼能代表臺灣？
蚵仔煎的魅力，
不在於精緻，
而在於它把幾種看似簡單的食材，
煎成了一種獨特的口感。
新鮮蚵仔的海味、
雞蛋的香氣、
地瓜粉形成的滑嫩外皮，
最後再淋上甜中帶鹹的醬汁，
一口下去，
就是夜市的完整畫面。
這種味道，
很難在其他國家找到替代品。

圓環邊，吃的是記憶感
圓環邊蚵仔煎，
沒有多餘的包裝，
也不刻意迎合潮流。
它留下來的原因很簡單
味道夠穩、節奏夠快、
讓人一吃就知道「對，就是這個」。
對旅客來說，
這是一家
不需要研究、不需要比較，就能安心點蚵仔煎的地方。

老臺北胃的帶路小提醒
第一次吃蚵仔煎，我會這樣建議：

1. 趁熱吃，口感最好
   
2. 不用急著加辣，先吃原味
3. 醬汁是靈魂，別急著把它拌掉

蚵仔煎不是細嚼慢嚥的料理，
它屬於人聲鼎沸、鍋鏟作響的夜市時刻。
站在人群裡，
吃著一盤熱騰騰的蚵仔煎，
你會很清楚地感受到
這，就是臺北的夜晚。

地址：103臺北市大同區寧夏路46號

電話：0225580198

菜單：https://oystera.com.tw/menu

第 6 家：阿淑清蒸肉圓｜第一次吃肉圓，就該從這裡開始

說到臺灣小吃，
 很多人腦中一定會出現「肉圓」兩個字。
但真正吃過之後才會發現，
 肉圓，從來不只有一種樣子。
在臺北，
 阿淑清蒸肉圓，
 就是我很常拿來介紹「清蒸派肉圓」的一家。

清蒸肉圓，和你想像的不一樣
不少旅客對肉圓的第一印象，
來自油炸版本，
外皮厚、口感重。
而阿淑的清蒸肉圓，
完全是另一個方向。
外皮晶瑩、滑嫩，
帶著自然的彈性，
不油、不膩，
一入口反而顯得清爽。
內餡扎實，
豬肉香氣清楚，
搭配特製醬汁，
味道層次簡單卻很乾淨。

為什麼我會推薦給第一次來臺北的旅客？
因為這顆肉圓，
不需要適應期。
它不刺激、不厚重，
即使是第一次嘗試臺灣小吃的人，
也能輕鬆接受。
對旅客來說，
這是一顆
「吃得懂、也記得住」的肉圓。

老臺北胃的帶路小提醒
第一次來阿淑，我會這樣吃：

1. 直接點一顆清蒸肉圓，吃原味
2. 醬汁先別全部拌開，邊吃邊調整
3. 放慢速度，感受外皮的口感變化

這不是夜市裡熱鬧喧囂的料理，
而是那種
安靜地展現臺灣小吃功夫的味道。
當你吃完這顆肉圓，
會更明白一件事
臺灣小吃的魅力，
往往藏在這些細節裡。

地址：242新北市新莊區復興路一段141號

電話：0229975505

第 7 家：胡記米粉湯｜一碗最貼近臺北早晨的味道

如果說前面幾樣小吃，
 是臺北的熱鬧與記憶，
 那麼米粉湯，
 就是這座城市最真實的日常。
而在臺北，
 胡記米粉湯，
 是很多人從小吃到大的存在。

為什麼米粉湯，這麼「臺北」？
米粉湯不是重口味料理，
它靠的不是刺激，
而是一碗清澈卻有深度的湯。
胡記的湯頭，
用豬骨慢慢熬出香氣，
喝起來清爽、不油，
卻能在喉嚨留下溫度。
米粉細軟，
吸附湯汁後入口順滑，
簡單到不能再簡單，
卻正是臺北人習以為常的早晨風景。

配菜，才是這一碗的靈魂延伸
在胡記吃米粉湯，
主角雖然是湯，
但真正讓人滿足的，
往往是那些小菜。
紅燒肉、豬內臟、燙青菜，
隨意點上幾樣，
湯一口、菜一口，
就是很多臺北人記憶中的早餐組合。
對旅客來說，
這是一種
不需要解釋，就能融入的臺北生活感。

老臺北胃的帶路小提醒
第一次來胡記，我會這樣建議：

1. 一定要點米粉湯，湯先喝
2. 再配 1～2 樣小菜，體驗會完整很多
3. 這一餐適合慢慢吃，不用趕

這不是為了觀光而存在的小吃，
而是一碗
每天準時出現在臺北人生活裡的湯。
當你坐在店裡，
聽著湯勺碰撞的聲音，
你會突然感覺到——
原來，臺北的早晨，
就是從這樣一碗米粉湯開始的。

地址：106臺北市大安區大安路一段9號1樓

電話：0227212120

第 8 家：藍家割包｜一口咬下的臺灣街頭記憶

如果要選一道
 外國旅客一看到就會好奇、吃完又會記住的小吃，
 割包，一定在名單裡。
而在臺北，
 藍家割包，
 就是我很放心帶旅客來認識這道經典的一站。

割包，為什麼被叫做「臺灣漢堡」？
割包的結構其實很簡單：
鬆軟的白饅頭、
燉得入味的滷五花肉、
酸菜、花生粉、香菜。
但真正迷人的，
是這些元素組合在一起時，
形成的層次感。
肉香、甜味、鹹味、清爽度，
在一口之間同時出現，
沒有誰搶戲，
卻彼此剛好。
這種平衡感，
正是臺灣小吃很迷人的地方。

藍家割包不是走浮誇路線，
它給人的感覺很直接
就是你期待中的割包樣子。
饅頭柔軟不乾，
五花肉肥瘦比例恰到好處，
入口即化卻不膩口，
花生粉的甜香收尾，
讓整體味道非常完整。
對第一次吃割包的旅客來說，
這是一個
不會出錯、也很容易愛上的版本。

老臺北胃的帶路小提醒
第一次吃藍家割包，我會這樣建議：

1. 直接點招牌割包，不要改配料
2. 如果有香菜，建議保留，味道會更完整
3. 趁熱吃，饅頭口感最好

割包不是精緻料理，
卻非常有記憶點。
站在街頭，
拿著一顆熱騰騰的割包，
邊走邊吃，
你會很清楚地感受到
這一口，就是臺灣的街頭生活。

地址：100臺北市中正區羅斯福路三段316巷8弄3號

電話：0223682060

菜單：https://instagram.com/lan_jia_gua_bao?utm_medium=copy_link

第 9 家：御品元冰火湯圓｜臺北夜晚最溫柔的一碗甜

吃了一整天的臺北小吃，
 到了這個時候，
 胃其實已經差不多滿了。
但只要天氣一涼，
 或夜色慢慢降下來，
 你還是會想找一碗——
 不是為了吃飽，而是為了舒服的甜點。
這時候，我通常會帶你來 御品元冰火湯圓。

為什麼叫「冰火」？這碗湯圓的關鍵就在這裡
御品元最有特色的地方，
就在於它的「冰火交錯」。
熱騰騰的湯圓，
外皮軟糯、內餡濃香，
搭配冰涼清甜的桂花蜜湯，
一口下去，
溫度在嘴裡交替出現。
不是衝突，
而是一種很細膩的平衡。
這樣的吃法，
也正是臺灣甜點很擅長的地方——
不張揚，但很有記憶點。

這是一碗，會讓人慢下來的甜點
和夜市裡熱鬧的甜品不同，
御品元的冰火湯圓，
更像是一個讓人停下腳步的存在。
你會發現，
坐在這裡吃湯圓的人，
說話聲都會不自覺地變小。
對旅客來說，
這不只是吃甜點，
而是一個
把白天的熱鬧慢慢收進回憶裡的時刻。

老臺北胃的帶路小提醒
第一次吃御品元，我會這樣建議：

1. 點招牌冰火湯圓，體驗完整特色
2. 先單吃湯圓，再搭配湯一起吃
3. 放慢速度，這一碗不適合趕時間

這不是為了拍照而存在的甜點，
而是一碗
會讓你記得「那天晚上在臺北，很舒服」的湯圓。

地址：106臺北市大安區通化街39巷50弄31號

電話：0955861816

菜單：https://instagram.com/lan_jia_gua_bao

第 10 家：頃刻間綠豆沙牛奶專賣店｜把臺北的味道，留在最後一口清甜

走到這一站，
 其實已經不需要再吃什麼大份量的東西了。
這時候，
 最適合的，
 是一杯不吵鬧、不張揚，
 卻會默默留在記憶裡的飲品。
頃刻間綠豆沙牛奶，
 就是我很常用來替一天畫下句點的選擇。

綠豆沙牛奶，為什麼這麼「臺灣」？
在臺灣，
飲料不只是解渴，
而是一種生活節奏。
綠豆沙牛奶看起來簡單，
但真正好喝的版本，
靠的是火候、比例，
還有耐心。
頃刻間的綠豆沙，
口感細緻、不粗顆，
甜度自然、不膩口，
牛奶的加入，
讓整杯變得柔順而溫和。
這不是衝擊味蕾的飲料，
而是一種
喝完之後，會覺得剛剛那一刻很舒服的甜。

為什麼我會用它當作最後一站？
因為它很臺北。
你可以外帶，
邊走邊喝；
也可以站在店門口，
慢慢把杯子喝空。
沒有儀式感，
卻很真實。
對旅客來說，
這杯綠豆沙牛奶，
就像是把今天吃過的所有味道，
溫柔地整理好，
帶走。

老臺北胃的帶路小提醒
第一次喝頃刻間，我會這樣建議：

1. 直接點招牌綠豆沙牛奶
   
2. 正常甜就很剛好，不用特別調整
3. 找個角落慢慢喝，別急著趕路

這一杯，
不會讓你驚呼，
卻會在回程的路上，
突然想起來。
原來，臺北的味道，是這樣結束一天的。

地址：111臺北市士林區小北街1號

電話：0228818619

菜單：https://instagram.com/chill_out_moment?igshid=YmMyMTA2M2Y=

如果只有 3 天的自助旅行在臺北，怎麼吃這 10 家？

第一次來臺北，
時間有限、胃容量也有限，
與其每一家都趕，不如照著節奏吃。
這份 3 天小吃路線，
是老臺北胃會帶朋友實際走的版本：
不爆走、不硬塞，
讓你每天都吃得剛剛好。

臺北 3 天小吃推薦行程表（老臺北胃版本）

天數	時段	店家名稱	小吃類型
Day 1	午餐	饌堂－黑金滷肉飯（雙連店）	滷肉飯
Day 1	下午	阿淑清蒸肉圓	肉圓
Day 1	晚餐	富宏牛肉麵	牛肉麵
Day 1	宵夜	胖老闆誠意肉粥	粥品
Day 2	早餐	胡記米粉湯	米粉湯
Day 2	下午	藍家割包	割包
Day 2	晚上	士林夜市－吉彖皮蛋涼麵	涼麵
Day 2	夜市	圓環邊蚵仔煎	蚵仔煎
Day 3	下午	御品元冰火湯圓	甜點
Day 3	收尾	頃刻間綠豆沙牛奶專賣店	飲品

雖然每個小吃的地點都有一點距離，但是你也知道，好吃的小吃，是值得你花一點時間前往品嘗
老臺北胃的小提醒

1. 不需要每一家都點到最滿
2. 留一點餘裕，才會想再回來
3. 臺北小吃的魅力，不在於吃多少，而在於記住了什麼味道
   

當你照著這 3 天走完，
你會發現，
臺北不是靠一兩道名菜被記住的，
而是靠這些看似日常、卻很真實的小吃。
下次再來，老臺北胃再帶你吃更深的那一輪。

老臺北胃帶路｜這 10 口，就是我心中的臺北

寫到這裡，
 其實已經不是在推薦哪一家小吃了。
而是在回頭看，
 這座城市，是怎麼用食物陪著人生活的。
滷肉飯、牛肉麵、肉粥、米粉湯，
 不是為了成為觀光名單而存在，
 而是每天默默出現在臺北人的日子裡。
夜市裡的蚵仔煎、涼麵、割包，
 熱鬧、吵雜、節奏很快，
 卻也正是臺北最真實的樣子。
而最後那碗湯圓、那杯綠豆沙牛奶，
 則是在一天結束時，
 替味蕾留下一個溫柔的句點。

如果你問我，
「這 10 家是不是臺北最好吃的小吃？」
我會說，
它們不一定是排行榜第一名，
卻是我真的會帶朋友去吃的版本。
因為它們吃得到：

1. 臺北人的日常
2. 巷弄裡的熟悉感
3. 不需要解釋，就能被理解的味道

如果你是第一次來臺北，
跟著這份清單走，
你不一定會吃得最飽，
但你一定會記得——
臺北，是什麼味道。
而如果有一天，
你又再回到這座城市，
走進熟悉的街口、
看到冒著熱氣的小攤，
你也會開始懂得，
為什麼老臺北胃，
總是記得這些看似平凡的滋味。
因為，真正留在心裡的，
從來不是吃過多少，
而是哪一口，讓你想起臺北。

胖老闆誠意肉粥真的好吃嗎？

走完這 10 家，

你可能會發現一件事阿淑清蒸肉圓真的有誠意嗎？

臺北的小吃，其實不急著被你記住。

它們就安靜地存在在街角、夜市、轉彎處，御品元冰火湯圓吃起來順口嗎？

等你有一天，再回到這座城市。阿淑清蒸肉圓冬天適合吃嗎？

如果你是第一次來臺北，胖老闆誠意肉粥新手友善嗎？

希望這份「老臺北胃帶路」的清單，

能幫你少一點猶豫、多一點安心。

不用擔心踩雷，饌堂－黑金滷肉飯（雙連店）不點會後悔嗎？

也不用為了排行而奔波，胡記米粉湯第一次適合嗎？

只要照著節奏走，

你就會吃到屬於自己的臺北味道。

而如果你已經來過臺北，

那更希望這篇文章，胡記米粉湯怎麼點比較好？

能帶你走進那些

你可能錯過、卻一直都在的日常小吃。

因為真正迷人的旅行，

從來不是把清單全部打勾，

而是某一天，

你突然想起那碗飯、那口湯、那杯甜，胖老闆誠意肉粥值得排隊嗎？

然後在心裡對自己說一句：士林夜市－吉彖皮蛋涼麵需要特地跑一趟嗎？

「下次再去臺北，還想再吃一次。」

把這篇文章存起來、分享給一起旅行的人，

或是在規劃行程時，再回來看看。

讓味道，成為你認識臺北的方式。

下一次來臺北，

別急著走遠。

老臺北胃，阿淑清蒸肉圓辣的推薦嗎？

會一直在這些地方，

等你再回來。

Digital reconstructions of human neurons overlaid on a slice of brain tissue donated by a brain surgery patient. Allen Institute researchers are able to capture electrical information from these live human neurons, as well as their 3D shape and gene expression, through a technique known as Patch-seq. This image shows several different types of human neurons in the medial temporal gyrus of the neocortex, the outermost shell of the mammalian brain. Credit: Allen Institute Hundreds of neuroscientists built a ‘parts list’ of the motor cortex, laying groundwork to map the whole brain and better understand brain diseases. Before you read any further, bring your hand to your forehead. It probably didn’t feel like much, but that simple kind of motion required the concerted effort of millions of different neurons in several regions of your brain, followed by signals sent at 200 mph (320 kph) from your brain to your spinal cord and then to the muscles that contracted to move your arm. At the cellular level, that quick motion is a highly complicated process and, like most things that involve the human brain, scientists don’t fully understand how it all comes together. Now, for the first time, the neurons and other cells involved in a region of the human, mouse, and monkey brains that control movement have been mapped in exquisite detail. Its creators, a large consortium of neuroscientists brought together by the National Institutes of Health’s Brain Research Through Advancing Innovative Neurotechnologies® (BRAIN) Initiative, say this brain atlas will pave the way for mapping the entire mammalian brain as well as a better understanding of mysterious brain diseases — including those that attack the neurons that control movement, like amyotrophic lateral sclerosis, or ALS. The atlas is described in a special package of 17 articles published today (October 6, 2021) in the journal Nature, including a single flagship paper that describes the entire atlas. Complete, brain-wide reconstructions of several different types of mouse neurons in 3D. A new study led by researchers at the Allen Institute and Southeast University in Nanjing, China, captured the detailed 3D shapes of more than 1,700 individual neurons in the mouse brain, the largest dataset of its kind to date. Studies like this will help neuroscientists piece together detailed views of neural circuits. Each color represents a different individual neuron. Credit: Allen Institute “In a human brain, there are more than 160 billion cells. Our brain has more than 20 times more cells than there are people in this world,” said Hongkui Zeng, Ph.D., Executive Vice President and Director of the Allen Institute for Brain Science, a division of the Allen Institute, and lead investigator on several BRAIN Initiative-funded studies. “To understand how a system works, you need to first build a parts list. Then you have to understand what each part is doing and put the pieces together to understand how the whole system works. That’s what we’re doing with the brain.” The massive BRAIN Initiative-funded collaboration involved dozens of research teams around the country who worked together to complete a cell-by-cell atlas of the primary motor cortex, a part of the mammalian brain that controls movement. Combining more than a dozen different techniques to define brain “cell types” across three different species of mammals, the resulting open-access data collection is by far the most comprehensive and detailed map of any part of the mammalian brain ever released. The researchers classified the millions of neurons and other kinds of brain cells present in the motor cortex into many different cell-type categories — the actual number of different brain cell types in this region depends on how they are being measured, but ranges from several dozen to more than 100. Scientists at the Allen Institute are studying human neurons that appear to be highly specialized as compared to their rodent counterparts. One of these newly described neuron types, the CARM1P1 neuron, sends long-range connections in the brain and may be selectively vulnerable in Alzheimer’s disease. Credit: Allen Institute The researchers picked the primary motor cortex in part because it’s similar across all mammalian species — while humans, monkeys and mice have many differences between our brains, the way we control movement is very similar — and because it’s representative of the neocortex, the outermost shell of the mammalian brain that not only integrates sensory and motor information but also gives rise to our complex cognitive functions. This completed atlas is one large step in the effort to create a catalog or census of all brain cell types through the BRAIN Initiative Cell Census Network, or BICCN. The NIH launched the BICCN in 2017, awarding nine collaborative network grants, three of which are led by Allen Institute for Brain Science researchers. Like a population census, the cell census aims to catalog all different types of brain cells, their properties, their relative proportions, and their physical addresses to get a picture of the cell populations that together form our brains. Knowing the “normal” brain’s cellular makeup is a key step to understanding what goes wrong in disease. “If we really want to understand how the brain works, we have to get down to its fundamental unit. And that is the cell,” said Ed Lein, Ph.D., Senior Investigator at the Allen Institute for Brain Science and lead investigator on several BRAIN Initiative studies focused on the human brain. “This is also clinically important because cells are the locus of disease. By understanding which cells are vulnerable in different brain diseases, we can better understand and ultimately treat the diseases themselves. The hope with these studies is that by making this fundamental classification of cell types, we can lay the groundwork for understanding the cellular basis of disease.” The atlas’s creators used several different methods to measure a variety of cellular properties to define a cell type by correlating and integrating these properties, which include the complete set of genes a cell switches on; a cell’s “epigenetic” landscape, which defines how genes are regulated; cells’ 3D shapes; their electrical properties; and how they connect to other cells. The single-cell gene expression and epigenetic data were especially important as the researchers were able to use these data to integrate all the other kinds of cell-type data, creating a common framework to classify cell types and compare them within and between species. The studies required not only collaboration among researchers to design and execute the experiments, but also coordination and public sharing of the data that resulted from the atlas project and other projects under the BICCN. The Brain Cell Data Center, or BCDC, is headquartered at the Allen Institute. The data center, led by Allen Institute for Brain Science Investigator Michael Hawrylycz, Ph.D., helps to organize the BICCN consortium and provides a single point of access to the study’s data-archiving centers across the country. “One of our many limitations in developing effective therapies for human brain disorders is that we just don’t know enough about which cells and connections are being affected by a particular disease, and therefore can’t pinpoint with precision what and where we need to target,” said John Ngai, Ph.D., Director of the NIH BRAIN Initiative. “The Allen Institute has played an important role in coordinating the large amounts of data produced by the BRAIN cell census project that provide detailed information about the types of cells that make up the brain and their properties. This information will ultimately enable the development of new therapies for neurologic and neuropsychiatric diseases.” Scientists at the Allen Institute for Brain Science played a role in nine of the 17 published studies and led or co-led six of them. The four primary Allen Institute-led studies explored: How cell types in the primary motor cortex compare across mice, humans, and marmoset monkeys. The research team found that most motor cortex brain cell types have similar counterparts across all three species, with species-specific differences at the level of proportions of cells, their shapes and electrical properties, and individual genes that are switched on and off. For example, humans have about twice as many excitatory neurons as inhibitory neurons in this region of the brain, while mice have five times as many. The researchers also delved into the famous Betz cells, enormous neurons that project to the spinal cord that exist in us, monkeys and many other larger mammals, and captured the first known electrical recordings from human Betz cells, which degenerate in ALS. Mice have evolutionarily related neurons based on shared genetic programs, but their shapes and electrical properties are very different from those in humans. A broader analysis of brain cell types in the human brain, looking at the second and third layers of the 6-layered neocortex. These layers, and the neocortex overall, are much larger and contain a larger diversity of cells in humans and other primates as compared to rodents. Allen Institute researchers used a three-prong technique known as Patch-seq to measure the electrical properties, genes and the 3D shapes of several kinds of neurons in these layers in tissue samples donated by brain surgery patients. The study characterizes these neurons in living human tissues and demonstrates an increased diversity of the types of neurons specialized to communicate between different regions of the human cortex, including delving into a specialized type of human neuron that is especially vulnerable in Alzheimer’s disease. The largest collection to date of complete brain-wide reconstructions of more than 1,700 different neurons in the mouse brain. This form of 3D neuron-tracing is extensive and complicated due to the cells’ lengthy and delicate axons and dendrites, but it yields important information about the long-distance connections different neuron types make through their axon arbors reaching faraway brain regions. Allen Institute researchers find that these neurons’ axon arbors show extremely diverse patterns, some with just a few focused branches while others spread across large areas. For example, some neurons in the structure known as the claustrum send axon arbors in a crown-like fashion around the entire circumference of the neocortex. Characteristic connection patterns like these are a critical attribute used to help classify a brain cell type. The cellular makeup of the mouse primary motor cortex, sorting approximately 500,000 neurons and other brain cells into cell-type categories based on the suite of genes each cell switches on (the “transcriptome”) as well as the gene-regulatory modifications on a cell’s chromosomes (the “epigenome”). Using a range of techniques, Allen Institute researchers and their collaborators generated seven types of transcriptomic and two types of epigenomic datasets, then developed computational and statistical methods to integrate these datasets into shared “evolutionary tree” of cell types. The study led to the discovery of thousands of marker genes and other DNA sequences specific for each of these cell types. Reference: “A multimodal cell census and atlas of the mammalian primary motor cortex BRAIN Initiative Cell Census Network (BICCN)” by BRAIN Initiative Cell Census Network (BICCN), 6 October 2021, Nature. DOI: 10.1038/s41586-021-03950-0 This research was supported by several awards from the National Institutes of Health, including award numbers U19MH114830, U01MH114812, U01MH105982, R01EY023173, and U24MH114827 to Allen Institute for Brain Science researchers. The content is solely the responsibility of the authors and does not necessarily represent the official views of NIH and its subsidiary institutes.

Moon phases. New Evidence Links Moon Phases to Human Sleep Cycles For centuries, humans have blamed the moon for our moods, accidents, and even natural disasters. But new research indicates that our planet’s celestial companion impacts something else entirely — our sleep. In a paper published on January 27, 2021, in Science Advances, scientists at the University of Washington, the National University of Quilmes in Argentina, and Yale University report that sleep cycles in people oscillate during the 29.5-day lunar cycle: In the days leading up to a full moon, people go to sleep later in the evening and sleep for shorter periods of time. The research team, led by UW professor of biology Horacio de la Iglesia, observed these variations in both the time of sleep onset and the duration of sleep in urban and rural settings — from Indigenous communities in northern Argentina to college students in Seattle, a city of more than 750,000. They saw the oscillations regardless of an individual’s access to electricity, though the variations are less pronounced in individuals living in urban environments. The pattern’s ubiquity may indicate that our natural circadian rhythms are somehow synchronized with — or entrained to — the phases of the lunar cycle. Sleep Disruption Peaks Before a Full Moon “We see a clear lunar modulation of sleep, with sleep decreasing and a later onset of sleep in the days preceding a full moon,” said de la Iglesia. “And although the effect is more robust in communities without access to electricity, the effect is present in communities with electricity, including undergraduates at the University of Washington.” Using wrist monitors, the team tracked sleep patterns among 98 individuals living in three Toba-Qom Indigenous communities in the Argentine province of Formosa. The communities differed in their access to electricity during the study period: One rural community had no electricity access, a second rural community had only limited access to electricity — such as a single source of artificial light in dwellings — while a third community was located in an urban setting and had full access to electricity. For nearly three-quarters of the Toba-Qom participants, researchers collected sleep data for one to two whole lunar cycles. Past studies by de la Iglesia’s team and other research groups have shown that access to electricity impacts sleep, which the researchers also saw in their study: Toba-Qom in the urban community went to bed later and slept less than rural participants with limited or no access to electricity. But study participants in all three communities also showed the same sleep oscillations as the moon progressed through its 29.5-day cycle. Depending on the community, the total amount of sleep varied across the lunar cycle by an average of 46 to 58 minutes, and bedtimes seesawed by around 30 minutes. For all three communities, on average, people had the latest bedtimes and the shortest amount of sleep in the nights three to five days leading up to a full moon. New research shows that on nights before a full moon, people sleep less and go to bed later on average. The pattern’s ubiquity, which was observed in urban and rural settings, may indicate that our natural circadian rhythms are somehow synchronized with the phases of the lunar cycle. Credit: Rebecca Gourley/University of Washington When they discovered this pattern among the Toba-Qom participants, the team analyzed sleep-monitor data from 464 Seattle-area college students that had been collected for a separate study. They found the same oscillations. Moonlight as an Ancient Signal for Activity The team confirmed that the evenings leading up to the full moon — when participants slept the least and went to bed the latest — have more natural light available after dusk: The waxing moon is increasingly brighter as it progresses toward a full moon, and generally rises in the late afternoon or early evening, placing it high in the sky during the evening after sunset. The latter half of the full moon phase and waning moons also give off significant light, but in the middle of the night, since the moon rises so late in the evening at those points in the lunar cycle. “We hypothesize that the patterns we observed are an innate adaptation that allowed our ancestors to take advantage of this natural source of evening light that occurred at a specific time during the lunar cycle,” said lead author Leandro Casiraghi, a UW postdoctoral researcher in the Department of Biology. Image of the moon. Credit: University of Washington Whether the moon affects our sleep has been a controversial issue among scientists. Some studies hint at lunar effects only to be contradicted by others. De la Iglesia and Casiraghi believe this study showed a clear pattern in part because the team employed wrist monitors to collect sleep data, as opposed to user-reported sleep diaries or other methods. More importantly, they tracked individuals across lunar cycles, which helped filter out some of the “noise” in data caused by individual variations in sleep patterns and major differences in sleep patterns between people with and without access to electricity. Artificial Light Mirrors Lunar Sleep Patterns These lunar effects may also explain why access to electricity causes such pronounced changes to our sleep patterns, de la Iglesia added. “In general, artificial light disrupts our innate circadian clocks in specific ways: It makes us go to sleep later in the evening; it makes us sleep less. But generally, we don’t use artificial light to ‘advance’ the morning, at least not willingly. Those are the same patterns we observed here with the phases of the moon,” said de la Iglesia. “At certain times of the month, the moon is a significant source of light in the evenings, and that would have been clearly evident to our ancestors thousands of years ago,” said Casiraghi. The Gravitational Semilunar Hypothesis The team also found a second, “semilunar” oscillation of sleep patterns in the Toba-Qom communities, which seemed to modulate the main lunar rhythm with a 15-day cycle around the new and full moon phases. This semilunar effect was smaller and only noticeable in the two rural Toba-Qom communities. Future studies would have to confirm this semilunar effect, which may suggest that these lunar rhythms are due to effects other than from light, such as the moon’s maximal gravitational “tug” on the Earth at the new and full moons, according to Casiraghi. Regardless, the lunar effect the team discovered will impact sleep research moving forward, the researchers said. “In general, there has been a lot of suspicion on the idea that the phases of the moon could affect a behavior such as sleep — even though in urban settings with high amounts of light pollution, you may not know what the moon phase is unless you go outside or look out the window,” said Casiraghi. “Future research should focus on how: Is it acting through our innate circadian clock? Or other signals that affect the timing of sleep? There is a lot to understand about this effect.” Reference: “Moonstruck sleep: Synchronization of human sleep with the moon cycle under field conditions” by Leandro Casiraghi, Ignacio Spiousas, Gideon P. Dunster, Kaitlyn McGlothlen, Eduardo Fernández-Duque, Claudia Valeggia and Horacio O. de la Iglesia, 27 January 2021, Science Advances. DOI: 10.1126/sciadv.abe0465 Co-authors are Ignacio Spiousas at the National University of Quilmes; former UW researchers Gideon Dunster and Kaitlyn McGlothlen; and Eduardo Fernández-Duque and Claudia Valeggia at Yale University. The research was funded by the National Science Foundation and the Leakey Foundation.

Recent animal studies have highlighted the crucial role of brain cells called astrocytes in sleep regulation. New research shows that activating these cells can keep mice awake for extended periods without making them sleepier later. This finding might lead to interventions that reduce the negative impacts of prolonged wakefulness, potentially benefiting shift workers, first responders, and military personnel. The Role of Astrocytes in Sleep Regulation New animal research suggests that little-studied brain cells known as astrocytes are major players in controlling sleep needs and may someday help humans go without sleep for longer without negative consequences such as mental fatigue and impaired physical health. Published in the Journal of Neuroscience, the study found that activating these cells kept mice awake for hours when they would normally be resting, without making them any sleepier. “Extended wakefulness normally increases sleep time and intensity, but what we saw in this study was that despite hours of added wakefulness these mice did not differ from well-rested controls in terms of how long and how intensely they slept,” said senior author Marcos Frank, a neuroscientist and professor at the Washington State University Elson S. Floyd College of Medicine. “This opens up the possibility that we might someday have interventions that could target astrocytes to mitigate the negative consequences of prolonged wakefulness.” Potential Applications for Shift Workers and Military Frank envisioned that might include medications that could be used to improve the productivity, safety, and health of shift workers and others who work long or odd hours, such as first responders and military personnel. Sleep loss and mistimed sleep have been shown to impact a variety of key processes, including attention, cognition, learning, memory, metabolism, and immune function. Astrocytes: More Than Just Brain “Glue” Astrocytes are types of non-neuronal cells that interact with neurons, brain cells that transmit easily measured electrical signals from the brain to other parts of the body. Previously thought of as merely the “glue” that holds the brain together, astrocytes have recently been shown to play an active role in various behaviors and processes through a much more subtle and difficult-to-measure process known as calcium signaling. This includes a previous WSU study that showed that suppressing astrocyte calcium signaling throughout the brain resulted in mice building up less sleep need after sleep deprivation. In this study, the researchers looked specifically at astrocytes in the basal forebrain, a brain region known to play a critical role in determining time spent asleep and awake as well as sleep needs. Using chemogenetics—a method to control and study signaling pathways within brain cells—they activated these astrocytes and found that this resulted in mice staying awake for 6 hours or more during their normal sleep period. What’s more, the researchers did not see subsequent changes in sleep time or sleep intensity in response to the added wakefulness, as would be expected. “Our findings suggest that our need for sleep isn’t just a function of prior wake time but is also driven by these long-ignored non-neuronal cells,” said first author Ashley Ingiosi, an assistant professor of neuroscience at Ohio State University who conducted the study while working as a postdoctoral research associate in Frank’s lab at WSU. “We can now start to pinpoint how astrocytes interact with neurons to trigger this response and how they drive the expression and regulation of sleep in different parts of the brain.” Future Research Directions Next, the researchers plan to conduct behavioral tests in mice to determine how activating basal forebrain astrocytes to induce wakefulness might impact other processes besides sleep needs, such as attention, cognition, learning, memory, metabolism, and immune function. To get at least some indication of the potential impact on attention and cognition, they looked at EEG markers of those two processes in this study and found them to be similar to those seen in well-rested controls. Reference: “Activation of Basal Forebrain Astrocytes Induces Wakefulness without Compensatory Changes in Sleep Drive” by Ashley M. Ingiosi, Christopher R. Hayworth and Marcos G. Frank, 8 August 2023, Journal of Neuroscience. DOI: 10.1523/JNEUROSCI.0163-23.2023 The study was funded by the National Institute of Neurological Disorders and Stroke, the National Institute of Mental Health, and the National Institute of Neurological Disorders and Stroke.

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