這個東西真的不錯

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）

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。

胖老闆誠意肉粥辣的推薦嗎？

走完這 10 家，

你可能會發現一件事御品元冰火湯圓有名是真的嗎？

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，御品元冰火湯圓當正餐適合嗎？

等你有一天，再回到這座城市。御品元冰火湯圓推薦必點嗎？

如果你是第一次來臺北，士林夜市－吉彖皮蛋涼麵口味會太清淡嗎？

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，頃刻間綠豆沙牛奶專賣店值得一試嗎？

也不用為了排行而奔波，士林夜市－吉彖皮蛋涼麵年輕人會喜歡嗎？

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，圓環邊蚵仔煎值得排隊嗎？

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，圓環邊蚵仔煎適合第一次吃嗎？

然後在心裡對自己說一句：士林夜市－吉彖皮蛋涼麵有名是真的嗎？

「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，饌堂－黑金滷肉飯（雙連店）有必要排隊嗎？

會一直在這些地方，

等你再回來。

An illustration of Tiktaalik, an ancient species among one of the first to transition to land. A new study reveals clues how the first animals on land evolved to eat. Credit: Illustration by Kalliopi Monoyios Advanced Imaging, Modern Species Provide New Insights Into Behavior of Tiktaalik Roseae New research out of the University of Chicago has found evidence that the lobe-finned fish species Tiktaalik roseae was capable of both biting and suction during feeding, similar to modern-day fish called gars. Scientists had been curious how the first animals on land evolved to eat, because most water-dwellers use suction to pull in food—which doesn’t work on land. The new results, published in the Proceedings of the National Academy of Sciences, provide evidence that bite-based feeding originally evolved in aquatic species and was later adapted for use on land. Life at the Water-to-Land Transition T. roseae, a creature whose flat skull is reminiscent of an alligator, is a species that lived “right at the cusp of the transition from life in water to life on land,” said senior author Neil Shubin, the Robert R. Bensley Distinguished Service Professor of Organismal Biology and Anatomy at UChicago. Studying its fossilized remains can provide new insights into how key traits for life on land originally evolved. “Water is different from air, being much denser and more viscous,” said Justin Lemberg, a postdoctoral researcher at UChicago and first author of the study. “This would have created unique problems for animals that were moving out of water and onto land for the very first time, including challenges in locomotion, reproduction, maintaining homeostasis and sensory processing and, of course, feeding. If you can’t feed yourself on land, how can you colonize it?” High-speed video of a baby alligator gar using cranial kinesis, suction, and biting to capture prey (similar to the feeding strategy proposed for Tiktaalik roseae). Credit: Video courtesy of Justin Lemberg (University of Chicago). Most aquatic vertebrate species use suction feeding to help pull prey into their mouths. To create suction, many species of fish can expand their skulls laterally to expand their mouths and produce negative pressure. This movement of the skull bones relative to one another is called cranial kinesis. “Suction feeding is ineffective on land, because it no longer works from a distance and it’s hard to create the pressure seal needed to draw something in,” said Lemberg. “So terrestrial vertebrates had to turn to other methods to capture prey. But the fossil evidence for how this happened is ambiguous, much more so than the transition from fin to limb. We wanted to look specifically at the sutures in the T. roseae skull, where the bones fit together, to see if they could tell us how the skull was being used.” The research team used advanced new computed tomography (CT) analysis to conduct a detailed examination of the morphology of the T. roseae skull. This allowed them to identify key new traits that had not been seen with other techniques, including sliding joints that would have allowed for the necessary cranial kinesis for the animal to expand its skull laterally to create suction. “We discovered Tiktaalik in 2004 and at the time, prepared it with the classical methods, removing rock from the fossil grain by grain,” said Shubin. “By the time Justin joined the project, we had access to this CT scanning technology, which lets us see the skull in 3D, taking each part out individually to see its shape and motion. Using the CT analysis transformed how we were able to think about the skull.” Side-by-side comparison of Tiktaalik (top) and alligator gar (bottom) showing similarly shaped snouts that may suggest convergence in feeding strategies. Credit: Image courtesy of Justin Lemberg (University of Chicago) Investigators noted distinct similarities between T. roseae and earlier work analyzing the skulls of alligator gar, a “living fossil” species previously thought to only use lateral snapping motions to capture prey. In a 2019 study, Lemberg et. al. found that gar use lateral snapping and suction synergistically while feeding, thanks to unique sliding joints in their skulls that help create suction while biting. Long-Standing Evolutionary Innovation These similarities led the researchers to believe that T. roseae may have fed in the same way, indicating that this adaptation likely evolved long ago, before animals ever colonized land. Disarticulated CT-based model of the skull of the alligator gar, showing joints between functional regions of the skull responsible for cranial kinesis. Credit: Image courtesy of Justin Lemberg (University of Chicago) “The thing that really stuns me is that every innovation, every invention used by tetrapods on land, originally appeared in some form in fish, including lungs, appendages, and now, feeding,” said Shubin.  Beyond teaching us about the evolution of our distant, fishy ancestors, better understanding of the biology and behavior of creatures like T. roseae can provide new insights into our own anatomy and development. Personal Connections to Ancient Evolution “The neat thing about the water-to-land transition is that it’s deeply personal to us,” said Lemberg. “How did we get to where we are now, and what are some of the evolutionary quirks we’ve adapted to get here?” Disarticulated CT-based model of the skull of Tiktaalik roseae, showing joints between functional regions of the skull responsible for cranial kinesis. Credit: Image courtesy of Justin Lemberg (University of Chicago) Case in point: Lemberg pointed out that when analyzing the range of motion for the T. roseae skull, the three bones that appear to have moved the most are the bones that would eventually become incorporated into the mammalian middle ear. “Those three bones in Tiktaalik are what we use to hear sound,” Lemberg said. “A little bit of cranial kinesis that’s maintained in modern mammals!” The other author was Edward B. Daeschler of Drexel University. Reference: “The feeding system of Tiktaalik roseae: an intermediate between suction feeding and biting” by Justin B. Lemberg, Edward B. Daeschler, and Neil H. Shubin, 1 February 2021, Proceedings of the National Academy of Sciences. DOI: 10.1073/pnas.2016421118 Funding: Two anonymous donors; the Academy of Natural Sciences of Philadelphia; the Brinson Foundation; the Putnam Expeditionary Fund (Harvard University); the University of Chicago; the National Geographic Society Committee for Research and Exploration; the National Science Foundation.

Color-coded nerve fiber orientations in a brain section obtained from scattered light imaging (SLI, left) and small-angle X-ray scattering (SAXS, right). Credit: TU Delft The Scattering Light Imaging (SLI) technique provides a cost-effective, high-resolution method to map neural connections in the brain. The technique, which involves analyzing light scattering patterns in thin brain slices, offers more detailed results than existing methods like dMRI, and is more accessible and faster than SAXS. Disentangling the complex nerve fiber network of the brain is becoming easily accessible with scattered light imaging (SLI): researchers in Delft, Jülich (Germany), and Stanford (USA) successfully combined light and X-ray scattering with MRI to discern nerve fiber trajectories, also in regions with highly entangled fibers. SLI revealed the trajectories at highest detail, while being significantly faster and cheaper than X-ray and MRI techniques. Such detailed mapping is essential for a better understanding of how nerve fibers are wired inside the brain. Pathways in the Brain The different areas in the brain are connected to each other via billions of nerve fibers. These connections are vital for proper brain function. The quest for a comprehensive map of all neural connections critically depends on imaging techniques that can disentangle these fibers, most of them only about a micrometer thin. Particularly challenging are regions with densely packed and highly interwoven nerve fibers. Miriam Menzel, Assistant Professor at the Department of Imaging Physics of TU Delft, developed the SLI technique to study such fiber constellations: “We shine light under different angles through hair-thin brain slices and analyze the resulting scattering patterns. We’re not taking a picture of neurons or synapses; we want to know how they are wired. This is important for understanding brain function and dysfunction.” More Accessible, Cheaper and Faster Small-angle X-ray scattering (SAXS) is an established method in material science to look into how different structures are organized with a synchrotron, while diffusion magnetic resonance imaging (dMRI) is an important technique in clinics for visualizing the three-dimensional nerve fiber network of the brain. “We have now shown that SLI data are consistent with those from SAXS and dMRI in the examined brain slices, but SLI provides higher resolution than dMRI and is more accessible, cheaper and faster than the other techniques. This is an important milestone,” Menzel says. “We can perform SLI measurements with a simple LED light source and camera in just a few seconds, requiring neither a multi-million synchrotron nor an MRI scanner. As a portable system, it could easily be set up in pathology laboratories to assist clinical research.” Microscopic Resolution Menzel has spent the past few years working on the SLI technique, first in Jülich and now in Delft. She also implemented it in Stanford, where her fellow researchers performed SAXS and dMRI measurements on brain samples also imaged with SLI. “Most imaging techniques struggle to discern individual pathways in dense brain structures containing many entangled or interwoven nerve fibers,” Menzel explains. “SLI provided fiber orientation maps with microscopic resolution in these dense regions.” Especially the two-dimensional (“in-plane”) fiber orientations were discerned with high precision. Next Steps “Being in Delft brings exciting opportunities to develop the technique further and work on new applications,” says Menzel. The team plans to also apply SLI to other types of fibers, such as muscle and collagen fibers, and to enlarge the tissue area that can be studied. The aim is to develop a small and portable system that can easily be deployed in other labs. “In the long term, we hope to apply the technique in clinics as well.” Reference: “Using light and X-ray scattering to untangle complex neuronal orientations and validate diffusion MRI” by Miriam Menzel, David Gräßel, Ivan Rajkovic, Michael M Zeineh and Marios Georgiadis, 11 May 2023, eLife. DOI: 10.7554/eLife.84024

Researchers deciphered the structure of an ion channel from the rod cells of the eye (shown in blue) while it interacts with the protein calmodulin (purple). This interaction is important to the function not only of the ion channel in the eye, but also of ion channels in other parts of the body such as the heart. Credit: Paul Scherrer Institute / Dina Schuster Exciting new findings shed light on the interaction between the protein calmodulin and an ion channel in the eye, potentially unlocking the secret behind our eyes’ exceptional sensitivity to low light conditions. Utilizing cryo-electron microscopy and mass spectrometry, a team of researchers from PSI has successfully unraveled the structure of an ion channel in the eye as it interacts with the protein calmodulin – a puzzle that has stumped scientists for 30 years. This interaction could explain how our eyes can achieve such remarkable sensitivity to dim light. The findings have been published in the journal PNAS. When you gaze at the bright screen of your phone or computer, the ion channels in your eyes react to the light by closing. This action marks the culmination of a biochemical chain reaction initiated by light exposure. As a result, calcium ions can no longer traverse the channels situated in the cell membrane, which leads to the transformation of the biochemical signal into an electrical one. This signal then travels through the nervous system, ultimately reaching your brain for processing. The same process occurs when you stand outside at night and looks up at the sky. Now, the rod cells perform this trick. These are the cells that make our eyes sensitive to low levels of light, enabling us to look at the night sky and detect just a few photons of light from a distant star. We take this for granted, but this is a remarkable feat. A team led by PSI scientist, Jacopo Marino, has now improved our understanding of how a tiny protein called calmodulin helps to achieve this, by interacting with ion channels in the rod cells. Calmodulin is a calcium sensor. It enables the cell to respond to calcium fluctuations – one of the cell’s universal means of communication. The team, a collaboration between groups at PSI, ETH Zurich, and University of Bonn, has illuminated for the first time the three-dimensional structure of the rod cyclic nucleotide-gated (CNG) ion channel as calmodulin binds. An Important Function for Calmodulin in the Eye One year ago, the researchers succeeded in deciphering the structure of this same ion channel, found in the rod cells of a cow retina and identical to the ion channel found in the rod cells of our eyes. Rod CNG consists of four subunits, a structure shared with many other ion channels. Yet a peculiarity of the channel is that three subunits – known as subunit A – are identical, whilst a fourth – subunit B – is different. Scientists have known for a long time that this subunit binds calmodulin. Throughout the animal kingdom, this feature is found. Yet, the exact nature of its role has remained unclear. “If something is conserved through evolution, it’s a very strong indicator that it’s important in some way,” explains Marino. “We knew that calmodulin modulates the activity of the channel through subunit B, but which kind of structural changes were occurring has been a big mystery for about thirty years, essentially because people were unable to solve the structure of the ion channel.” Now, the researchers can provide a three-dimensional view of what is really happening. Through a combination of cryo-electron microscopy and mass spectrometry, they could observe that as calmodulin binds, the ion channel becomes a bit more compact. The researchers believe that this is nature’s way of holding the channels closed. What would the purpose of this be? “We think it’s a way to reduce spontaneous channel openings that would cause background noise so that our eyes can be sensitive to dim light,” says Marino. Mass Spectrometry Helps Researchers Solve a Wriggly Structure Obtaining the structure of calmodulin and the ion channel binding was not easy. The interaction between calmodulin and Rod CNG occurs in a highly flexible region of the channel, where it is free to swing about. In cryo-electron microscopy, this makes it very difficult to obtain high-resolution structural information. Here, Marino offers an analogy, “Imagine you have a room of people dancing. You take a photo and want to work out from this what the human body shape is. You might be able to work out what a head looks like, but with limbs waving all over the place the legs and arms will be blurred.” It was thanks to a chance meeting, that the team could pin down this wriggly structure. Ph.D. student Dina Schuster heard a presentation of Marino. “We were ready to publish based on the cryo-electron microscopy data alone, which left much of the interaction ambiguous, when Dina approached me and said ‘I think I can help you’“, he remembers. Schuster is developing novel mass spectrometry-based strategies to study protein interactions. These techniques use enzymes to chop proteins into pieces, either in native conditions within parts of the retinal membrane or when chemically crosslinked. The protein fragments, some of which are joined together, are identified by mass spectrometry. This reveals information on which parts of the protein were close together in three-dimensional space – equivalent to piecing together a 3D jigsaw puzzle. “These techniques enabled us to narrow down some of the possibilities that were ambiguous with cryo-electron microscopy,” explains Schuster, who is joint first author of the publication together with PhD student, Diane Barret. From the Wonder of Vision to Implications in Human Health Calmodulin regulates ion channels not only in the eye but throughout the body, controlling electrical signals that are essential to the correct functioning of diverse muscles and organs. In recent years, it has become apparent that when this interaction goes wrong due to mutations in the calmodulin gene, there can be severe health implications, such as cardiac failure: something that is not yet fully understood. As well as helping our understanding of a most fundamental wonder – how we can see the stars – the findings of this study, and methods used, may aid our understanding of the interaction of calmodulin with ion channels in other parts of the body. Reference: “Structural basis of calmodulin modulation of the rod cyclic nucleotide-gated channel” by Diane C. A. Barret, Dina Schuster, Matthew J. Rodrigues, Alexander Leitner, Paola Picotti, Gebhard F. X. Schertler, U. Benjamin Kaupp, Volodymyr M. Korkhov and Jacopo Marino, 3 April 2023, Proceedings of the National Academy of Sciences. DOI: 10.1073/pnas.2300309120

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饌堂－黑金滷肉飯（雙連店）值得專程去吃嗎？ 》台北大揭密｜10家美食名店全盤解析圓環邊蚵仔煎值得吃嗎？ 》台北夜市高分美食推薦｜10間絕對不踩雷士林夜市－吉彖皮蛋涼麵會踩雷嗎？ 》台北小吃美食餐廳怎麼挑？10家人氣店幫你選